ZOOV111/ZOOX111

Class notes
Prof Ronel Nel
2021
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Content: Diversity of Life

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Introduction brought about by human activities, resulting in a far

greater rate of extinction than ever experienced
Living organisms display a bewildering and truly
before. Up to a hundred species are thought to
remarkable range of sizes, shapes, structures and
become extinct every day! This is called the
life histories. Some are so small they can be seen
Anthropocene or the 6th extinction. Species are lost
only under an electron microscope (bacteria) others
due to an increased need for natural resources to
weigh many tons (elephants and blue whales).
feed and house a growing human population.
Some, like the largest trees (Sequoia), may live for a
thousand years or more, others for no more than a With a population at nearly 7.5 billion people,
few days (diatoms). Some obtain their energy natural environments are destroyed to develop
directly from sunlight (plants or autotrophs), some economic hubs, cities, and harbours, or forests and
from the products of decay (scavengers and other natural habitats are transformed to
decomposers), while others eat plants and other agricultural land for food production, or
organisms (heterotrophs) like predators or (over)exploited, fished and mined for natural
parasites. Some organisms are highly mobile so resources. The rate of species extinction and habitat
they can fly thousands of kilometres, or creep destruction is of great concern, as we know that the
around underground, and many are permanently presence of many different species is essential to the
attached to the substratum (sedentary). Moreover, maintenance of tolerable conditions on planet earth.
organisms have adapted to virtually every The species richness of 8.5 million extant
environment on earth, from hot springs to the polar species is only an estimate because many species
ice, from the deepest oceans to the highest remain unknown or undescribed whereas some
mountains, and from sun-baked deserts to the total have gone extinct before they were described. Of
darkness of caves. these known species alive today, nearly three
It is estimated that there are at least 8.5 million million are animal, with insects being the most
different species of organisms alive on earth today diverse with about 800 000 known species. There
of which only about one-and-a-half million have are only 4000 species of mammals. Southern Africa
been named. A far larger number are now extinct. In is particularly well endowed with biological
fact, throughout evolutionary history, extinction has diversity - we have > (more than) 700 species of
been the norm rather than the exception. It is butterflies in the region and > 900 species of birds;
brought about by such factors as changes in the flora of the Cape Peninsula has more plant
environmental conditions, or a failure to compete species, and a higher concentration of threatened
successfully with other organisms for resources, in species, than any area of similar size in the world. If
short supply such as food or space. Extinction has you collected all the aquatic organisms you could
eliminated not only individual species but also find in False Bay, near Cape Town, you could end up
whole groups of organisms (like dinosaurs). A major with 2 500 species of algae, animals and
problem today is the drastic environmental changes microorganisms.
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CLASSIFICATION OF LIVING ORGANISMS birds in the same group, and that fishes and whales
are placed together, which clearly won't do.
Such numbers and diversity are bewildering and
Nevertheless, the principle was sound, that
confusing; our minds are such that we are compelled
organisms can be placed in large groups using
to try to bring order to such phenomena by
specific criteria and these groups then progressively
classifying them into some kind of system. We are all
subdivided different, but equally objective, criteria.
stamp collectors at heart, grouping similar objects
It is the basis of modern classification.
into categories and labelling them. Such groupings
may, in fact, have practical value. For example, Classification, as well as the naming of organisms

primitive peoples tended to group organisms into (nomenclature), lacked clear rules until the middle

those that were good to eat and those that were of the 18th century, when two circumstances

poisonous or distasteful, those that were beneficial combined to make the systematic study of

in treating the sick and those that ill people should organisms more rewarding as a discipline and more

avoid, and so on. Even such an essentially subjective exact as a science. One of these was the work of

classification demands experience and Carolus Linnaeus and his invention of binomial

observational powers. nomenclature (see below). The other was the

growing acceptance of the concept of evolution.
The first people to attempt a fully objective
Before the acceptance of evolution, classification
classification of plants and animals were the ancient
was just a matter of convenience; but if all species
Greeks, a people who were keen on pursuing
had indeed evolved from common ancestors, then
knowledge for its own sake. They knew more
they were all related to one another in varying, and
species than anyone before and went out of their
potentially measurable, degrees. Clearly, a good
way to collect information about plants and animals
system of classification should reflect these
in other lands. Aristotle (385-322 B.C.) was the most
relationships. This concept gave impetus to the
avid collector of biological information in the
study of biology and resulted in revision after
ancient world and speculated widely on natural
revision of the classificatory system over the past
phenomena, including the most appropriate system
two centuries.
of objective classification.

In the system of animal classification favoured by TAXONOMY

the ancient Greeks, a most important criterion was
The classification of organisms, or taxonomy, is
the colour of the blood; they divided the whole
based, as we have seen, on a hierarchical system of
animal kingdom into species with red blood and
groups within groups, with each group ranked at a
those without red blood. They then subdivided these
particular level. Each group is called a taxon and its
groups according to mode of locomotion - walking,
level of ranking in the system its category. At the
creeping, running, flying or swimming. This was a
time of Linnaeus (1707-1778), only three categories
naive system compared with present-day
were in common usage: the species, the genus
classification and if we apply it we find bats and
(containing species resembling one another in
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appearance), and the kingdom (a much higher to the group of species making up the genus - for
category). Linnaeus and subsequent biologists example, one may talk of Drosophila (the genus of
added more categories between genus and kingdom, fruit fly) without adding a species name. Both genus
so that today species are grouped into genera, and species names are commonly derived from
genera into families, families into orders, orders Latin or have Latin endings. By convention they are
into classes and classes into phyla or divisions. The printed in italics or written underlined. In the
sequence (from highest to lowest): phylum - class - taxonomic literature, the genus and species names
order- family - genus - species, is easily are followed by the name of the person who first
remembered by the mnemonic "Please Carry On For described that species and often by the date of that
Goodness Sake". The terms phylum and division are description. So, for example, Antidorcas marsupialis
exactly equivalent; for no good reason, phylum is Sundevall 1847 is the springbok as described by
used for animals and protozoans, while the term Sundevall in 1847.
division is generally used for prokaryotes, algae,
But why not just use common names that
fungi and plants. Although the categories given
everyone can understand? The first problem is that
above remain the basic ones, biologists have found
most species do not have common names. Worse,
it convenient to insert additional categories such as
where a common name does exist, it is often used for
suborder or superfamily. Ideally the categories
several species instead of just one. Thirdly, many
should reflect evolutionary relationships, species
well-known organisms have several common
within a single genus being very closely related to
names; for example, the names gnu and wildebeest
one another, while those in the same phylum (or
refer to the same animal, while Fiscal Shrike,
division) but in different classes, would be only very
butcher bird and Jackie hangman are common
distantly related.
names which relate to the same species of bird.
The system of binomial nomenclature Moreover, different languages have different
(doublenames) devised by Linnaeus are still in use common names for the same organism. It is
today. The name of a species consists of two words, therefore essential to have a naming system, which
the name of the genus (with a capital letter) is specific, unambiguous, not dependent on language
followed by the specific name (with a small letter). and universally recognised by all biologists, and
In order to avoid confusion, every genus must have informative.
a different name, but the species epithet need only
The International Commission on Zoological
be unique within the genus. Thus, names such as
Nomenclature (ICZN) is the authority on species
Callorhynchus capensis (the St Joseph shark),
names for animals. The Code (i.e. the rules) used as
Dichistius capensis (the galjoen) and Janolus capensis
well as an up-to-date list of officially accepted
(a sea slug) are all perfectly valid but only when both
animal species names can be downloaded from
genus and species names are given. ("Capensis"
http://www.iczn.org.
simply means "of the Cape".) On the other hand, the
name of the genus can stand alone if one is referring
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THE SPECIES CONCEPT clones of identical cells. So although botanists and

microbiologists use the term "species", this is a
Species is the Latin word for kind. So basically,
convenient category rather than representing real
different species are simply different kinds of
entities.
organisms. A much more rigorous definition is
needed, however, if we are to proceed scientifically. A practical difficulty with a species concept based

For one thing, forms that look very different may in on genetic isolation is that many taxonomists have

fact belong to the same species; for example, males never seen their organisms alive, much less have any

and females of a species may differ considerably in information as to their interbreeding potential.

appearance (this is called sexual dimorphism), and Classic taxonomy was based entirely on the

larval or juvenile stages may look totally different organism's structure (morphology) and it is only in

from the adult. Conversely, two different species recent decades that other features, including

may resemble one another so closely that only an physiology and behaviour, have been taken into

expert or genetics can tell them apart. account. A major advance has been the development
of techniques in molecular biology which allow DNA
A scientific definition of species, accepted by
sequencing and analysis of proteins. The greater the
zoologists, is that they are groups of actually or
similarity of the genes (DNA) and consequently of
potentially interbreeding natural populations,
the proteins, the more closely are the two organisms
which are reproductively isolated from other
presumed to be related. This holds true at all
groups. This definition makes sense because if two
taxonomic levels, including the species level.
groups exchange genes they will not develop or
maintain any characteristic differences between A good example of what can be done using DNA

them. One slight problem with the definition is that similarity is the story of the quagga. The quagga was

closely-related animal species will sometimes mate once abundant in the Cape but was hunted to

with one another - horses and donkeys, for example extinction by the mid-1880's. Although clearly

- and produce offspring. However, such offspring related to the zebra in appearance, it was regarded

(mules) are usually sterile; they cannot reproduce as a separate species. In the 1980's a scraping of cells

and so their combinations of genes are not passed was obtained from a museum specimen's skin.

on to future generations. However, this is not always Remarkably, enough intact DNA was still present to

the case. allow sequencing. This was then compared with

zebra DNA, leading to the conclusion that the quagga
While the above definition works well for animal
had been a zebra subspecies (or race), rather than a
species, it is not so good for plants and
distinct species. A selective breeding programme is
microorganisms. Many plants can reproduce
now under way, breeding zebras with quagga-like
asexually and can also form fertile hybrids with
characteristics.
another species. Bacteria display several types of
gene exchange, while many unicellular eukaryotes
Make sure that you can differentiate among
(e.g. Amoeba) reproduce by cell division, producing
the terms species, populations and communities
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and how we define it in terms of species richness because, like animals, they do not undertake
and species diversity which incorporates a photosynthesis. They have cell walls, but these
measure of abundance. consist of substances such as chitin, not
polysaccharides. These (and other) features suggest
THE QUESTION OF KINGDOMS that the fungi arose from Protista separately from
Until quite recently, all living things were plants and animals. They are thus accorded their
assigned to only two kingdoms - plants and animals. own kingdom. We now recognise at least five
This however does not reflect true evolutionary kingdoms of organisms.
relationships. Prokaryote micro-organisms (such
as bacteria), which lack a nucleus and have a THE ROLE OF TAXONOMY
singular, circular DNA ribbon not bonded to Taxonomy once thought of, as being dull and
protein, differ from both plants and animals to an pedestrian by ecologists, physiologists and
even greater extent than do plants from animals. biochemists, is in fact fundamental to the whole of
They must therefore have their own kingdom - the biology. It is not only essential to the study of
Monera. evolutionary relationships (phylogeny) but is also

Unicellular (or acellular) eukaryotes are basic to all ecology, including such practical matters

obviously closer to plants and animals than are the as biological assessment, conservation and parks

Monera and generally have some plant-like or management. An ecologist cannot possibly identify

animal-like characteristics but cellular organisation with certainty all the organisms he encounters and

and modes of reproduction are sufficiently different leans on expert taxonomists for this purpose.

to warrant giving them their own kingdom - the Experimental work would also be of no value if the

Protista. Some simple multicellular organisms are scientists could not be certain what species they

also included in this kingdom, because they were studying. More recently ecologists also count

resemble unicellular species more than advanced on genetics expertise to assist with species

multicellular organisms. identifications and population delineations.

To the plant kingdom is allocated those truly

multicellular organisms, which obtain their energy
from photosynthesis, having chlorophyll
contained in chloroplasts (autotrophic). They also
have cell walls with a matrix of polysaccharide (e.g.
cellulose). In contrast, organisms in the animal
kingdom have no chlorophyll (or chloroplasts) and
typically obtain their energy and nutritional
requirements from eating other organisms
(heterotrophic). Their cells do not have cell walls.
This leaves the fungi without a taxonomic home
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microscope, they show considerable diversity (>64K

named species) and often have complex life

PROTOSTOMIA histories. They are also extremophiles living in

habitats from the tropics to the Antarctic, in fresh
water, in the sea and in the soil, and their cysts are
THEME: SINGLE-CELLED ORGANISMS
often borne on the wind. A great many are parasites
responsible for a number of diseases, which affect
KINGDOM PROTISTA:
humans, but many have formed beneficial
(OR PROTOZOA AND CHROMISTA) relationships with other organisms or are of

Previously single-celled animals were allocated economic importance. They are represented by a

to the group Protozoa and single-celled large number of phyla, and due to the complexity of

photosynthesizers to the group Protophyta or Algae. the phylogeny, we will concentrate on studying four

However, protozoans are no longer considered to be different body shapes of Protists, which resemble

animals by most biologists. Instead algae and different animal cell types. We will use locomotion

protozoans are now grouped into the Kingdom as a means to discriminate among four different

Protista. Both Algae and Protozoa no longer have types of Protists.

any formal taxonomic status except as informal,

convenient groupings. The advent of the molecular 1. The Flagellates:
age scrambled our classic arrangement of single- Phylum Euglenozoa
celled organisms which was mostly based on form The name Flagellate is derived from the feature
and function. The most important characteristic of that protozoa belonging to these taxa are typically
Protista though is that they are single cells propelled by one (flagellum) or more hair-like
(unicellular), and each individual is surrounded by flagella. Flagellum means “whip” and the action of
a plasma membrane. Each individual also has at the flagellum is indeed sometimes whip-like,
least one nucleus surrounded by a nuclear although more commonly it propels the organism by
membrane. They are thus eukaryote. In this means of either an undulating or a corkscrew
respect, they resemble multicellular plants and motion. Although superficially resembling the
animals but differ from the bacteria which lack a flagellum of a bacterium, the protozoan flagellum
true nucleus and thus prokaryote. Protists may be has a quite different structure. In eukaryotes,
unicellular but they are not simple. They are including protozoa, each flagellum has a number of
complete organisms with all the functions of microtubules running along its length internally.
complex, multicellular organisms. These microtubules are always arranged in the same
pattern viz. nine pairs of microtubules
How did the eukaryotic cell originate?
surrounding a central pair. Below the base of each
Although individual Protists are all very small, flagellum, in the cytoplasm, is a basal body
the vast majority visible only with the aid of a (kinetosome), which consists of nine groups of
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microtubules; here each group consists of three flagellates may then release nutrients to the termite,
microtubules with no central pair of microtubules. although the termite’s chief source of nutrition is
almost certainly the dead bodies of the flagellates
The flagellates belong to different phyla, but
themselves. The termite and flagellate are
generally can be separated based on being
completely dependent on one another for survival, a
autotroph or heterotroph. Autotrophs synthesize
relationship termed symbiosis.
their own food from substances available in the
environment through photosynthesis or In humans, flagellates from a variety of genera
chemosynthesis, whereas heterotrophs can’t and (plural of genus) cause illness such as Giardia
derive their food from other organisms. causing diarrhoea or Trichomonas causing sexually
transmitted diseases. These effects may be serious
Class Euglenoidea in young children or people which are already

These are flagellated protistans more closely immuno-compromised.

allied to plants than to animals, often possessing

chloroplasts and are thus capable of Phylum Dinoflagellata
photosynthesis (e.g. Euglena). Coloured flagellates occur abundantly in the sea;
among these are the dinoflagellates (“whirling
Class Trypanosomatidae whip”), many of which are spectacular in high

Although most flagellates are free living in fresh abundances. Noctiluca causes bioluminescence

water or the sea, a few are parasitic. By far the most turning waves into spectacular light shows at night

important genus of parasitic flagellates in Africa is whereas others cause red tides or harmful algals

Trypanosoma, a genus containing a number of blooms (HABs). The phenomenon of red tides, which

species, which cause diseases in humans or other occurs commonly around the South African coast

animals. The most serious of these is sleeping and sometimes kills vast numbers of marine

sickness, caused by Trypanosoma gambiense or T. animals, due to nerve poisons contained in the cell.

brucei. The vector for the parasite is the Tsetse fly.

Other common diseases include Leishmaniases 2. The Ciliates:
(“white leprosy” or “black fever”) caused by Phylum Ciliophora
Leishmania spp. As the name implies, the Ciliophora (cilia
Some flagellates (from different phyla) living in bearing) are distinct from other protistans in the
the alimentary canals of animals should be regarded possession of numerous cilia, which are used both
as commensal rather than parasitic. This is true, for for locomotion and to create feeding currents in the
example, of Trichonympha, a flagellate living in the water. Each cilium has the same structure as a
termite gut. The termite eats wood but cannot digest flagellum and is supported by the same
it; it simply chews the wood up into small enough configuration of microtubules. A cilium may, in fact,
pieces for it to be ingested by Trichonympha. The be regarded as nothing more than an extremely
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short flagellum; it thus becomes reasonable to cause various diseases in humans and animals
deduce the evolution of Ciliophora from the although they rely on an intermediate host as
flagellated protozoans. vector (animal that spreads and transmits a disease
or parasite). It is characteristic of the group that a
Ciliates have also evolved two types of nuclei –
single individual undergoes multiple fission to give
macronuclei and micronuclei. A typical ciliate has
rise to numerous smaller cells, or spores; hence the
one nucleus of each kind. The macronucleus is
former name of the phylum, Sporozoa. Although
essential for the carrying out of cellular functions,
there is a very large number of species, one has
whereas the micronucleus is concerned with
outweighed all others in terms of medical, economic
sexual reproduction. Only the micronucleus
and historical importance. This is the malarial
undergo meiosis; haploid nuclei are then exchanged
parasite, Plasmodium vivax.
between two individuals during a process known as
conjugation. After exchange, the micronuclei fuse, The malarial parasite, Plasmodium vivax
so that each individual again has a diploid nucleus.
Plasmodium is carried by the female Anopheles
The micronucleus then divides mitotically, one of
mosquito and is injected into the human blood
the new daughter cells giving rise to a new
stream when the mosquito bites the victim. (Only
macronucleus, which dissolved during conjugation.
female mosquitoes suck blood.) There are no
Paramoecium is the best-known genus of symptoms for about 12 days after the parasites have
Ciliophora and its distribution in fresh- water ponds been injected. During this time they have been
and dams is worldwide. Some forms of ciliates are carried in the blood to the liver, where they enter the
sessile (attached to the substratum) and show a liver cells and divide by multiple fission into spores.
reduction in the number of cilia. These are no longer At the end of the 12 day period, the spores burst out
habitually used for locomotion but are used mainly of the liver cells into the blood and each spore enters
to create feeding currents. Examples are Stentor and a red blood corpuscle. Inside the corpuscle the spore
Vorticella, both of which are common in South grows, becoming amoeboid as it does so, and then

Africa. assumes the shape of a ring. This is the “signet ring

stage” which doctors look for to confirm their
diagnosis of malaria. The parasite then again
3. The Sporozoans:
undergoes multiple fission until the whole corpuscle
Phylum Apicomplexa
is crammed with spores. The pressure of these
Members of the Apicomplexa (“twisted tip”) all
spores causes the corpuscle to burst, releasing both
have a definite shape, like the flagellates, but lack
the spores and their accumulated waste products
organelles concerned with locomotion. They almost
into the blood. As this event is more or less
certainly evolved from flagellates, but the flagellum
synchronised for all the infected corpuscles, the
was lost in the process. As they have no means of
patient receives a massive dose of toxins into the
moving about, it is hardly surprising that all are
blood stream, resulting in the first bout of malarial
parasitic, with complicated life histories. Most
fever. The released spores enter other red blood
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corpuscles or liver cells and the cycle is repeated Sarcodinia. However, it now realised that this is not
over and over again, leading to regular bouts of fever a monophyletic group an several clades are present.
alternating with periods of more or less normal
There can be little doubt that some of the
health. However, as the disease progresses, such a
amoeba-like organism evolved from flagellated
large proportion of the red corpuscles have been
protozoans, particularly as species intermediate
destroyed that the patient becomes chronically
between the two classes exist. Naeglaria is one
anaemic and weakens progressively.
example. In Trichonympha part of the body is
After some time, some of the spores develop into flagellate, part amoeboid; but in Naegleria the body
male and female gametocytes (i.e. precursors of sex is alternately flagellate and amoeboid. The adult
cells). These sexual stages develop no further form is a large multi-nucleated amoeba, lacking
unless they are sucked up by an Anopheles flagella and moving by means of amoeboid
mosquito. In the insect’s gut they become gametes movement. In response to changes in the
and male and female gametes fuse to form a zygote. environment, it may encyst and divide repeatedly
The zygote enters the wall of the insect’s gut and within the cyst. When conditions are right, these
forms a cyst. Inside the cyst it again undergoes daughter cells emerge from the cyst as small oval
multiple fission, eventually releasing a large flagellates with a fixed shaped. As they grow and
number of sporozoites, which find their way to the mature, they lose their flagella and become
mosquito’s salivary glands, ready for injection into amoeboid once more.
the next human host.
Naegleria is of some medical interest. Although
This type of life cycle, involving two hosts and an the organism normally lives a free existence in damp
alternation of sexual and asexual reproduction is soil, it sometimes finds its way into dams or
typical of Sporozoa. So is the relatively brief change swimming pools. If one gets into the nasal passages
to an amoeboid stage, indicating evolutionary links of a swimmer, it may bore into the brain
with the amoeba-like organisms. (braineating amoeba) and cause a fatal nervous
disease (primary amoebic meningoencephalitis) for

4. The Amoebas which there is no cure. A few such cases have been
reported in southern Africa.
Typical of the amoeba-like organisms is that,
unlike the flagellates and ciliates, the body shape is Another species of considerable interest to
not constant because pseudopodia are produced southern Africa is Entamoeba histolytica (Phylum
for feeding and locomotion. This form of Amoebozoa) and related species, which cause
locomotion, in which the cytoplasm flows or amoebic dysentery. They ulcerate the human
streams forward into pseudopodia, is known as intestine and may also cause liver abscesses.
amoeboid movement. In older classifications all the However, some amoebic rhizopods live quite
amoeba-like organisms were included under happily in the gut without causing damage to the
host.
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Amoeba proteus pseudopodia that extend through the holes of the

shell. Vast numbers of them occur in the oceans and
Amoeba is certainly the best-known example of
when they die their shells slowly sink to the bottom
and is often taught at school level as a typical
as marine “snow” and may cover the seabed to a
protozoan, which it is not. It is also frequently
considerable depth. Chalk cliffs (white cliffs of
referred to as a very simple life form, which is also
Dover) are typically composed of the shells of these
quite false. Amoeba are, in fact, highly specialised
organisms.
and complex. They are found in sluggish streams,
ponds and ditches in most parts of the world. The
shape is highly irregular and pseudopodia may be Phylum Radiolaria
extruded from any point of the body. There is thus The Radiolaria, like Foraminifera, are amoebalike
no symmetry of any kind and no front, or rear end. marine species; they construct complex little shells
They engulf food particles, which may be living or of silica (or strontium sulphate). The shells have
dead, by means of pseudopodia. This is known as numerous pores, through which axopodia (semi-
phagocytosis and results in food vacuoles, or permanent pseudopodia with central rod) are
phagosomes, which may be seen travelling around extruded. Like the Foraminifera, vast numbers occur
the cytoplasm while the food in them is being in some oceans, the tiny shells, or skeletons, sinking
digested. Undigested food is eventually voided to the to the bottom to form a “radiolarian ooze”.
outside of the body. A conspicuous feature of fresh- Finally, a few free-living flagellates are also
water amoebae is the contractile vacuole, which colonial. A colony is formed when cell division does
constantly expels the contents. not proceed to completion, so that the daughter cells
Although some sarcodinoids display sexual remain joined together by a short stalk. Such
reproduction, Amoeba reproduces only asexually, by colonial forms are not common, but they are of
mitotic cell division. Under adverse conditions it can interest with regards to possible evolutionary
encyst. The cyst wall is tough and strong and enables links; they present the first possible example and
the organism to resist desiccation and high mechanism for multicellular life.
temperatures. The cysts may be blown about by the
wind, thus distributing the species. Inside the cyst,
the amoeba divides repeatedly, so that when LOW LEVELS OF ORGANISATION
conditions are again favourable a number of small
amoebae emerges.
THEME: COLONIES OF SINGLE CELLS WORKING
COOPERATIVELY
Phylum Foraminifera
Choanoflagellates are an “in-between” group of
Foraminifera (hole bearing) are marine,
organisms with characteristics of both unicellular
amoeboid protozoa, which construct tiny shells
and multicellular forms. These organisms, like
of calcium carbonate. They produce slender
Salpingoeca rosetta can appear as single, flagellated
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collar cells (choanocytes) or as clusters or colonies flagellum. By beating their flagella, the choanocytes
of divided, but not separated groups of choanocytes create a water current, forcing a continuous stream
with cooperative functioning. As early as 1814, it of water through the sponge. Within the water are
was already suggested that choanoflagellates are a small food particles, which are captured by the
stepping-stone group, but recent genetic analysis choanocytes as the water passes through. This
confirmed this; a single genotype code for both method of obtaining food is known as filterfeeding,
single, and colonial string or star phenotype forms. as the animal is literally filtering its food out of the
water. Once filtered the water passes out through a
hole at the top of the sponge, known as the osculum.
KINGDOM ANIMALIA: The outer lining of the sponge is made up of

MULTICELLULAR ORGANISMS pinacocytes.

Another type of cell that sponges possess is

THEME: CELLULAR SPECIALIZATION amoebocytes. These are found between an outer

layer of dermal cells and the inner layer of
choanocytes. The amoebocytes have several
functions, including reproduction. All sponges are
Phylum Porifera (Sponges) hermaphrodite, and the amoebocytes produce

The Porifera, or sponges, are aquatic, both male and female gametes. The female gametes

filterfeeding animals, which consist of several are retained by the amoebocytes, while the male

types of cells. However, the cells do not form tissues gametes are released into the water via the osculum.

or organs and display a high degree of autonomy, so These male gametes then drift around until they are

that in some respects a sponge resembles a colony filtered out of the water by the choanocytes of

of cells rather than an integrated individual. This another sponge. The choanocytes recognise these as

impression is strengthened by the fact that sponges gametes, and they are not digested. Instead they are

possess high regenerative and reaggregative passed on to the amoebocytes to fertilise the egg

powers. Thus, young sponges can develop from cells, and a larva is produced. This larva swims for a

fragments of the parent sponge body, and if a sponge while before settling to give rise to a new sponge.

is forced through a sieve to separate the cells, the The other important function of the amoebocytes
cells will reaggregate to form a new sponge. is the formation of skeleton. In sponges, the skeleton

The most conspicuous feature of sponges is the is either made up of spicules or a collagenous

extensive system of pores and canals. It is these fibrous material known as spongin, or both. As the

pores, which give the phylum its name ("porus" = name suggests, spicules are spiky structures; these

pore, "fera" = bearing). Lining the inner surfaces of mesh together to give support to the sponge body.

these pores and canals are the collar cells or Because they are so spiky, they also aid in deterring

choanocytes, each one of which possesses a predators. As a result, few animals eat sponges, with
 14

the exception of some sea slugs, which graze on Class Calcarea

them.
Members of this class contain spicules composed
Sponges can be grouped based on the complexity of calcium carbonate. Sponges belonging to this
of the structure and canals. The simplest form are class are generally small (<10cm in height).
the asconoid sponges where the spongecoel is lined
with choanocytes. In the case of syconoid sponges, Class Demospongiae
the incurrent canals are lined with choanaocytes, This class contains more than 90% of all sponges.
whereas leuconoid sponges have got much greater Members of this class have silicon spicules which are
surface areas for filtration with both incurrent and arranged randomly, or no spicules at all. Those that
radial canals of which the radial canals are lined do not secrete spicules are supported by spongin or
with choanocytes. collagen fibres, or a mixture of both.

Class Hexactinellida
In these sponges the spicules are always
composed of silicon and have a six-rayed structure.
The spicules are usually arranged in complex
networks, and the sponges are cup- or vase-shaped.
These sponges are generally found in deep water
(200 - 2 000m), and may grow to a large size (10 –
100 cm high). An example is the beautiful Venus
Flower Basket Euplectella.

Fig. 1 The structure of sponges and the details about

the design of different sponges (Branch and
Branch 1988).

There are four classes of sponges, distinguished

on the basis of the chemical composition, shape and
distribution of skeletal materials.

Class Homoscleromorpha
Flat encrusting sponges with no spicules or
siliceous spicules.
 15

DIPLOBLASTIC: generations”. The two basic body forms are the

polyp and the medusa.
TWO-LAYERED ANIMALS
Polyps have an elongated, cylindrical body,

THEME: TISSUES SPECIALISATION AND which is attached to the substratum at the base.
ORGANIZATION INTO ECTODERM AND These cylindrical bodies are crowned by tentacles,
ENDODERM; RADIAL SYMMETRY which surround an opening on the upper surface.
The medusae are free-swimming and resemble
inverted polyps. They are umbrella-shaped, with a
fringe of tentacles surrounding the mouth, which
Phylum Cnidaria hangs downwards. However, unlike the polyp, most
The Cnidaria is a phylum of multicellular aquatic of the body of the animal is made up of mesogloea,
animals with a simple body structure. However, usually consisting of a non-cellular gelatinous
their cells show a higher level of cooperation than do substance.
those of the Porifera. Cnidarians are diploblastic,
Polyps are sessile and, although they are capable
their cells being organised into two well-defined
of some movement, they cannot move over large
layers, an outer ectoderm and an inner endoderm.
distances, and they can only reproduce asexually by
These two cell layers are separated by a jelly-like
budding. This makes the mixing of genes and
layer of mesogloea which provides support for the
dispersal to new areas difficult. Polyps thus bud off
animal. Cnidarians have a central, sac-like cavity, the
medusae, which are capable of both long distance
coelenteron or gastrovascular cavity, in which
movement and sexual reproduction. Separate
digestion takes place. There is a single opening
(dioecious) male and female medusae produce
surrounded by tentacles, which serves as both the
gametes that are fertilised in the water. The zygote
mouth and the anus. The cells of the ectoderm and
develops into a planula larva, which is ciliated and
endoderm show considerable differentiation; with
is thus capable of swimming. This planula larva lives
muscle cells and a nervous system and are
for a while in the plankton before settling to the
therefore capable of movement. However, they have
substratum and forming a polyp. This is the typical
no organs of respiration, circulation or excretion,
life cycle of cnidarians. However, in some groups of
and each individual cell must fulfil these functions
cnidarians, either the polyp or the medusa stage has
for itself. Stinging cells, or cnidoblasts, are
become reduced or entirely lost.
typically present on the ectoderm; hence the name
The Cnidaria are organised on a circular body
of the phylum.
plan and are thus radially symmetrical. This
The Cnidaria exhibit polymorphism (i.e. the
means that the animal can be cut into two equal
occurrence of structurally and functionally different
halves along any vertically plane, so long as this
types during the life cycle) and “alternation of
plane passes through the midpoint. These animals
thus have no left and right sides. Contrast this with
 16

bilaterally symmetrical animals, such as the Colonial hydroids consist of many polyps, which
flatworms in the following sections, which can only come together to form branched colonies joined by
be cut in one plane to produce equal halves. Radial a common "stem". In these, the individual polyps
symmetry may be an advantage to sessile animals are known as zooids, and may show division of
(which do not move about) in that animals can sense labour. One such colonial hydroid is Obelia, which
the environment equally well in all directions, and it is common around the coast of southern Africa. An
may collect food equally well from all directions. It Obelia colony is made up of two different types of
is, however, not an advantage for mobile organisms. polyps called hydranths and blastostyles. The
hydranths are the feeding polyps, and they thus
The Cnidaria are divided into five classes, which
have a mouth surrounded by tentacles with which
are distinguished predominantly by the relative
to capture food items. These hydranths are
importance of their polyp and medusa stages.
supported and protected by a cuplike structure

Class Hydrozoa called the hydrotheca. The second type of polyp, the
blastostyles, are concerned with reproduction.
The Hydrozoa include the hydroids and
They do not feed, relying on the hydranths for
bluebottles. They are small, usually colonial, and
nutrition. The transfer of nutrients from the
mostly marine. In the Hydrozoa, complete
hydranths to the blastostyles occurs through the
alternation of generations with both polyp and
coenosarc, a living stem-like structure connecting
medusa stages is the general rule, although there
the polyps of the colony. This coenosarc is covered
are some exceptions.
by a non-living protective layer known as the
Hydroids (order Hydroida) are a diverse group, perisarc. The lack of tentacles makes the
with more than 280 species recorded from the blastostyles vulnerable, and they are thus housed
marine environment around southern Africa. Most within a protective covering called the gonotheca.
hydroids are colonial, forming intricately The blastostyles produce the medusae by asexual
branched structures, which resemble miniature budding. When the medusoid buds are mature,
trees or feathers. they break away from the blastostyle, floating out

Solitary hydroids consist of a single polyp, and through the opening in the gonotheca.
resemble sea anemones (see later in this chapter).
A well-known example is Hydra, a freshwater
hydroid. Although Hydra is often used as an
example of the Hydrozoa, it is atypical of this class.
Unlike most other Hydrozoa, Hydra does not have a
medusa stage during its life cycle. Instead, the
polyps may reproduce either asexually by budding,
or sexually, developing both male and female
gonads within the same individual.
 17

Fig. 2 Life cycle of Obelia hydroid (from Branch and

Branch 1988).

The order Siphonophora are marine colonial

hydrozoa, and includes the blue-bottle Physalia,
which is common along the coast of southern Africa.
These colonies are made up of three different types
of polyps or zooids, each of which performs a
different function in the colony. Gonozooids, as the
name suggests, are responsible for reproduction,
while gasterozooids are involved in digestion. The
longest strands of polyp are made up by the
dactylozooids, which are responsible for both prey
capture and defence. The entire colony is supported
by a gas-filled float, which keeps the colony at the
Fig. 3 The bluebottle (Physalia physalis).
water surface.

Class Scyphozoa
The Scyphozoa are the jellyfish. In these
cnidarians the medusa is the dominant stage of the
life cycle, and the polyp is very small and solitary.
The body form of the scyphozoan medusa is similar
to the general medusoid form described earlier in
this chapter. The medusa is bell-shaped, and the
majority of the body is made up of mesogloea.
 18

However, unlike the mesogloea of the Hydrozoa, coelenteron is divided into a series of vertical
that of the Scyphozoa may be cellular in some cases. partitions by a number of sheet-like mesenteries.
The mouth hangs downwards, and is located at the These mesenteries may have several functions,
end of a tube known as the manubrium. Tentacles including increasing the surface area for digestion
hang from the outer edge of the medusa. Unlike the and aiding in supporting the animal.
hydrozoan medusae, those of the Scyphozoa contain
true muscle fibres, which allow them to swim. This
is achieved by rhythmic contractions of the bell,
which propels the animal through the water.
Statocysts located around the margin of the bell
allow the animal to orientate itself in the water.

The sexes are dioecious (separate) in jellyfish,

and fertilisation is external. In Aurelia, which is
common in southern African waters, the eggs and
sperm pass through the mouth and into the water. Fig. 5 An anemone (Pseudactinia flagellifera).

Sea anemones are solitary anthozoans, and the

polyp is often very large (one Australian species
grows to a diameter of 1 m!). Many anemones are
brightly coloured. In some cases, they warn
potential predators that they are harmful or toxic.

Corals are colonial anthozoans, all of which

produce some sort of exoskeleton. The soft corals,
as their name suggests, produce a "skeleton" which
is composed primarily of mesogloea, and is thus
Fig. 4 Life cycle of Jellyfish Chrysaora (from Branch
relatively soft and flexible. The sea fans, or
and Branch 1988).
gorgonians, belong to this group.

Class Anthozoa The second major group of anthozoan corals are

Sea anemones and corals comprise the class the hard, or stony, corals. The individual coral

Anthozoa. The Anthozoa differ from the Hydrozoa polyps are very similar to those of the anemones.

and Scyphozoa in that, not only is the polyp the Unlike anemones, the minute coral polyps form

dominant form, but the medusa stage has become dense colonies, the entire colony being housed

completely lost. Anthozoan polyps possess two within a calcium carbonate exoskeleton. It is these

unique features. They have a pharynx, which is a skeletons that form gigantic reefs (such as the Great

short tube, made up of ectoderm, extending from the Barrier Reef off the north-eastern coast of

mouth into the coelenteron. Below this pharynx, the Australia), which provide living space, food and
 19

shelter for more species of animals than are to be Class Cubozoa

found in any other marine habitat in the world.
These include the box-jellies which are medusoid
jellies with four sides to them. They were originally
placed with Schyphoza but the complexity of these
animals warranted their own class. They develop
directly from the polyp stage to the medusa stage
without going through an ephyra larval stage.
Furthermore, these are often called sea wasps as
they are of the most venomous animals in the world,
with venom powerful enough to kill a human. The

Fig. 6a Example of soft coral (Alcyonium sp.) (from largest species known is Chironex fleckeri.

Branch and Branch 1988).

Class Staurozoa

The staurozoa are a cross between the polyp and

medusa phase with the medusa being stalked and
attached to the substrate like a polyp. They are
commonly also known as “upside down” jellies
because the medusa is facing upwards with the
tentacles facing into the water column. The life cycle
is also different than the classes with alternation of
Fig. 6b Example of a colony of hard coral (from generations by the absence of cilia on the planula
Branch and Branch 1988).
larva. The larva thus crawl across the surface until it
finds suitable habitat, settle, and grow directly into
an adult medusa without going through
strobilation.

Phylum Ctenophora
The Ctenophora are the comb jellies or sea
gooseberries. They are similar to the Cnidaria in
Fig. 6c Surface detail of hard coral (from Branch and that they are diploblastic, radially symmetrical,
Branch 1988). and essentially marine. Indeed, the two used to be
placed in the same phylum. The most important
difference between them is the lack of stinging
 20

cells in the Ctenophora. Another difference is that Germinal layers and body cavities
the ctenophores have eight longitudinal ciliated
All metazoans (other than the sponges) are built on
bands running the length of the body, the beating of
remarkably similar lines, even though superficially
which results in forwards propulsion of the animal.
they may look very different from each other. Firstly,
These cilia beat in a wave-like fashion, creating
when any animal develops from egg to embryo, its
ripples of iridescent rainbow colours as they do so.
tissues differentiate early on into two germinal
layers: the ectoderm (which will later give rise to
the skin and its associated organs, and the nervous
system) and the endoderm (which later gives rise
to the inner lining of the gut and to associated organs
like the liver or digestive glands). In most animals, a
third layer soon develops between ectoderm and
endoderm. This is the mesoderm, which gives rise
to the muscular system, the blood vascular system
(if there is one) and the gonads.
Secondly, animals all fit into one of only five
fundamental 'architectural plans' (see Fig. 5 A-D),
and even these have a lot in common. The simplest
body plan is that of the diploblastic acoelomates -
the Cnidaria and Ctenophora (A). Between ectoderm
and endoderm is not mesoderm but a noncellular
jelly-like layer known as the mesogloea. The guts of
such animals are blind-ending (ie there is a mouth
but no anus) and there are no internal spaces where
organs can develop. (Remember that the lumen of
the gut is not truly 'inside' the animal, in that it is in
contact with the outside world.) The only way in
which an animal like a sea anemone can make use of
a hydrostatic skeleton is by holding water in its
gastrovascular cavity and tightly closing its mouth.
The bodies of triploblastic acoelomates (B) like the
Platyhelminthes and Nemertinea sandwich a layer of
loose, watery mesoderm between ectoderm and
endoderm. This means that organs like gonads can
develop within the body itself; it also means that the
 21

mesodermal tissue can act as a hydrostatic skeleton,

and that there is no body cavity.

The pseudocoelomate condition (C) is found in the

assemblage of organisms (previously known as
Aschelminthes) with the best-known phylum
Nematoda. A pseudocoel is a cavity between
mesoderm and endoderm. It is filled with fluid
and acts as a typical hydrostatic skeleton. Note that
the gut consists only of endoderm and so it cannot
be innervated, nor can it be muscular. Most don’t
have circular muscle and pseudocoelomate animals
thus rely on movements of the whole body (from
longitudinal muscles) to move food down the gut.
All other animals, including the annelids, molluscs,
arthropods, echinoderms and chordates, are
coelomate (D). By definition, this means that the
body cavity (the coelom) is lined throughout by
mesoderm. This has extraordinarily important
consequences, apart from the fact that the coelom
can act as a hydrostatic skeleton. Firstly, the
mesoderm lining the endoderm of the gut can form Fig. 7 Comparison among animal groups (taxa)
muscles. Thus, the gut can move independently of the based on number of germ layers and body cavity. A)
diploblastic animals with blind-ending gut.
rest of the body. This in turn means that relatively
Triploblastic taxa B) without coelom; C) with a
indigestible food like plant material can be churned pseudocoelom lined only partially with mesoderm;
up and digested slowly (non-coelomates are D) with coelom, which is lined completely with
mesoderm.
generally unable to eat plant food other than
phytoplankton). Secondly, the gut can become long
and coiled; thirdly, it can develop accessory glands
like salivary glands and livers. All of these features
lead to vastly more efficient digestion and the
possibility of feeding on living and dead plant
material. What is more, a muscular blood vascular
system (and heart) can develop in the mesenteries
that connect the outer and inner layers of mesoderm.
 22

TRIPLOBLASTIC: THREE-LAYERED symmetry (left and right sides of the body are

ANIMALS mirror images) is more suited to organisms that

move actively through the water. Other changes that
may be more fitted to an active life style include
THEME: TRIPLOBLASTICITY, BILATERAL
SYMMETRY, WITH THE FIRST ORGAN SYSTEMS dorso-ventral flattening giving dorsal (upper) and
AND CEPHALIZATION ventral (lower) surfaces of the body, and
development of anterior (front) and posterior
(back) ends. This last feature facilitates grouping of
The diploblastic body layout of the Cnidaria has sensory organs at the front, with neural cells
two cell layers: ectoderm on the outside and grouped together to form a primitive brain
endoderm lining the coelenteron. Between these is (cephalization). The anterior is where such animals
a jelly layer called the mesoglea (or mesogloea). In will most likely first encounter changes in the
triploblastic animals an extra layer of cells, the environment.
mesoderm, replaces the mesoglea. This triploblastic
Though they have an extra layer of cells and new
arrangement allows a greater division of work.
body layouts, flatworms and ribbon worms, in
Ectoderm is responsible for protecting the organism
common with the Cnidaria, are still acoelomate.
and produces the nervous tissue. Endoderm, in
Thus, they have no fluid-filled cavity (coelom) in
common with that in diploblasts, has digestive
the body between any of the cell layers.
function. The new mesoderm is sandwiched
Consequently, organs are surrounded by body cells
between the other two layers and isolated from the
and this limits their independence of movement.
environment. This protection provides the
The similarities and differences exhibited by
mesoderm with more controlled conditions, which
these groups suggest that flatworms and ribbon
allow development of diverse tissues that can fulfil
worms are stepping-stones of increasing complexity
increasingly complex requirements of the organism
between the Cnidaria and higher animals. There is a
more efficiently. In particular, this allows the
graduation from cnidarians to flatworms as there is
development of dedicated reproductive and
from flatworms to ribbon worms, which in turn lead
excretory systems.
on coherently to the nematodes in the next section.
This new level of complexity is apparent in
In addition to their place in this anatomical scheme,
Platyhelminthes (flatworms), Nemertea (ribbon
flatworms are also interesting because some species
worms) and all higher animals. With new complexity
cause serious disease in man and livestock.
comes opportunity for new lifestyles and
possibilities for experimenting with new body
arrangements. Radial symmetry is appropriate for
a floating organism such as a jellyfish as it travels
with the water current, or sessile organisms such as
anemones; these organisms must be sensitive to the
environment from all sides. In contrast, bilateral
 23

by pumping movements of the pharynx. In the gut, or

gastrovascular cavity, endoderm cells engulf food
particles by phagocytosis. The gut is blind-ended
TRIPLOBLASTIC ACOELOMATE
and so undigested food leaves through the mouth.
The nervous system is not much advanced from the
LOPHOTROCHOZOA nerve net of cnidarians but there is a concentration
(“CREST-BEARING” ANIMALS): of sensory cells (cerebral ganglia) and nervous tissue
at the anterior. The two nerve cords that run the
length of the body are not homologous with our own
spinal cord but may be precursors of our autonomic
Phylum Platyhelminthes
nervous system. Excretion is by means of flame cells
Platyhelminthes ("platy" = flat, "helminthes" = (protonephidia) that drain into ducts and out via an
worms: Greek) are the most primitive of the excretory pores. Reproduction is both sexually with
triploblastic animals. Four classes: Turbellaria, cross-fertilization as they are monoecious, or
Trematoda (flukes), Monogenea and Cestoda asexually by fission (splitting) or regeneration (grow
(tapeworms) are considered here. Of these, the back when broken).
turbellarians are mostly free living; the other three
groups are always parasitic and consequently show
a number of specialisations for the parasitic
lifestyle. Class Trematoda (Flukes)
Several thousand species have been described; all
Class Turbellaria are parasites. To aid them in this life style they are
Some 3000 turbellarian species are described. typically each armed with a pair of suckers. The
They are typically leaf-shaped worms, and although suckers were originally thought to be holes, hence
some may be 50 cm or more long, most are less than the name trematodes ("trema" = hole: Greek). Most
10 millimetres. Most turbellarians are free living but trematodes have an anterior mouth that opens into
Notoplana patellarum makes an interesting an oral sucker connected to a forked digestive tract,
exception: it lives commensally in the mantle folds of but in one large and important group (the
the limpet, Cymbula oculus, found on rocky shores of gasterostomes) the mouth is sited similarly to that in
the Cape Province. turbellarians. Since adult trematodes typically live in
the host’s digestive tract, they have a tough
Turbellarians have a variety of exceptional
digestion resistant covering called a tegument
shapes but a “typical” turbellarian has a head with
which, in contrast to turbellarians, lacks cilia.
eyespots and moves by gliding using cilia. There is no
mouth on the head but has a mouth and pharynx on Because of the low likelihood of transmission of
the midventral surface. They eat carrion or actively parasite stages from host to host, internal parasites
capture prey that is then held down and torn to put much effort into reproduction. Many eggs and
pieces with the pharynx. Detached bits are ingested other stages are required because so many get lost
 24

and never reach the intended next host. In sporocyst produces many similar daughter
consequence, a great proportion of the parasite body sporocysts, which find their way to the gonad and
is given over to reproductive organs. Indeed, on digestive gland of the snail. Depending on the fluke
maturity, some parasites become a bag of eggs. species, some mother sporocysts produce daughter
sporocysts and others produce more muscular,
The liver fluke and the blood fluke are of great
motile offspring called rediae (named after the
medical and veterinary importance but both in their
Italian microscopist Francesco Redi of the 17th
own way are atypical of the Trematoda.
Century). The redia, in contrast to the daughter
Nevertheless, their importance renders them
sporocyst, has a well-developed digestive system. In
preferable subjects for study rather than an
both rediae and daughter sporocysts, further asexual
anatomically typical but otherwise obscure fluke.
processes produce balls of cells that develop into the
tadpole-like cercaria. Many cercariae have tails and
are able to swim. Their sensory organs enable them
to detect a host and by using penetration apparatus
they pass through its skin. If in the final host, usually
a vertebrate, cercariae shed their tails and migrate in
the blood to the optimum site in the host. Here they
develop into adults and begin laying eggs. In some
trematodes the cercariae encyst; this can be on
vegetation, inside the next intermediate host
(usually an invertebrate or lower vertebrate) if there
is one in the life cycle, or even in the snail host. The
Fig. 8 General life cycle of a trematode (from cyst stage contains a metacercaria ("meta" = after
Hickman et al. 2012). or later: Greek). If it is ingested by the final host, the
cyst is digested away and the freed metacercaria
The liver fluke and the blood fluke described
develops into an adult fluke.
below have two-host life cycles; other trematodes
may have three hosts or more. The first intermediate Liver fluke (Fasciola hepatica)
host is usually a snail and the final or principal host Adults of the liver fluke can reach 25 millimetres
is a vertebrate. Eggs from the adult fluke escape to in length. This fluke is hermaphroditic, as are most
the environment where they hatch, releasing a trematodes, but in contrast with typical trematodes
ciliated free-swimming larva called a miracidium. the reproductive and digestive systems are
This minute larva, a fraction of a millimetre in length, branched. The life cycle follows that described above
then seeks a suitable snail host. It penetrates, loses except that the cercaria encysts on vegetation near
its cilia, and migrates in the snail’s circulatory watercourses and waits to be eaten by the final host
system to a suitable site. Here it develops into a (sheep, cow or man). In the final host it escapes from
simple bag-like mother sporocyst. By an asexual the cyst and migrates from the stomach to the liver
process, known as polyembryony, the mother
 25

via the bile duct were it develops into the adult. It from the human host into water hatch into
damages the bile duct walls leading to blood loss and miracidia, which seek and penetrate water snails.
reduction in bile flow. A bizarre and aberrant human Here they go through sporocyst stages to release
infection called halzoun has been reported from cercariae, which penetrate the final host’s skin
some areas in the Middle East where raw liver is (human). The cercaria loses its tail and the infective
customarily eaten. Should this contain liver flukes stage, called a schistosomula, eventually arrives in
they are able to attach themselves temporarily to the the blood vessels of the liver where it matures and
pharynx causing pain and distress. finds a mate to complete the cycle.

Blood fluke (Schistosoma)

Class Monogenea
This is the causative organism of the disease
The monogeneans are typically ectoparasites
bilharzia. Some 300 million people suffer from blood
previously lumped with the Trematoda but may be
fluke infections. The name Schistosoma is derived
more closely related to the Cestoda. Monogeans are
from schisto: split and soma: body (in Greek). The
ectoparasites on aquatic animals like frogs and
split is the groove in the male (10 mm long) in which
turtles where they attached to the gills or surfaces by
the female (15 mm long) permanently resides.
means of an opisthaptor (a large sucker with
Separate sexes such as in Schistosoma are atypical
hooks). Like most other flatworms, these animals are
of the trematodes. Hermaphroditism is more
monoecious, fertilized eggs hatch into a ciliated
common because the transmission from host to host
larva and after a brief free-living phase attach to a
is uncertain and the likelihood of individual flukes of
single host where they complete their life history.
the opposite sex meeting in the host is consequently
These animals are generally not life threatening but
low. Schistosoma compensates at least in part by
is a problem animal in aquaculture conditions where
meeting and pairing up in the liver, using it as a
hosts live in high densities.
clearing house, before migrating, joined together, to
the blood vessels of the bladder.
Class Cestoda
The female lays a large number of eggs each day.
Adult cestodes are known as tapeworms (the
Each egg has a spine and releases enzymes that
name cestode is derived from kestos = strap: Greek).
soften the host tissue. The pulse of the blood works
Several thousand species have been described and
the spine through the vessel wall and the eggs pass
all are parasitic. Some are of great economic and
into the bladder. This causes much blood loss to the
medical importance. Depending on species they vary
urine and the host frequently suffers anaemia with
from a few millimetres to about 30 metres. They are
consequent fatigue and lethargy. The eggs also lodge
perhaps the most specialised of the worm parasites;
in capillaries of the liver and lungs where they kill
an indication of this is their absent digestive
tissues and cause fibrous growths. The infection also
system. Who needs one when nutrients can be
has a more sinister aspect. The irritant effect of the
absorbed through the body surface? A further
spine and enzymes can induce development of
consequence of a parasitic life style is that much
urinary system cancers. Eggs that manage to escape
body space is given over to reproductive organs.
 26

General life cycle of tapeworms pass out with the faeces. When these are ingested by
the second host, they hatch in the gut as hexacanth
The adult tapeworm usually lives in the digestive
larvae or oncospheres ("hexacanth" = six spines;
tract of a vertebrate host and it is essentially an
"onco" = tumour: Greek, because the oncosphere
eggproducing machine with an anchor. The anchor is
swells tumour-like into a cysticercus). The
called the scolex; it is armed with suckers and a
hexacanth penetrates the gut wall and travels
rostellum of hooks. The egg producing part, the
through the blood to muscle and nervous tissue of
strobilus, is a chain of flattened, and usually
the second host (normally a pig or cow according to
rectangular, segments called proglottids (or
worm species). Here it develops into a cysticercus
proglottides). These are joined to the scolex by the
10-12 millimetres in diameter with a scolex that is
neck. New proglottids develop from the neck and
pushed into the cyst and inverted like the finger of a
each new proglottids remains joined to the next. As
glove. When man ingests the cysticercus, the scolex
older proglottids are pushed away by growth of new
everts and fixes the larva to the gut wall. Proglottids
ones at the neck, a flat tape-like chain of proglottids
develop from the neck and a new tapeworm adult
is so produced. Each proglottid has both male and
completes the cycle.
female reproductive organs (monoecious) but the
male system matures first. In the older proglottids Eggs of Taenia solium are also infective to man
the female system is predominant. By folding of the and, as in the pig, they develop into cysticerci in the
strobilus in the intestine of the host the active male tissues. Cysticerci can occur in human muscle tissue,
proglottids are brought into contact with the older heart, brain, spinal cord and eyes. This stage is
proglottids containing mature female organs and clearly more harmful than infection with the adult
fertilization of the eggs can occur. The proglottids worm. A significant proportion of human hosts with
then becomes a bag of eggs. Eventually it detaches adult Taenia solium also harbour cysticerci in their
from the end of the tape and may disintegrate to tissues. It appears that proglottids may be
release the eggs or it may pass intact out of the host regurgitated from the small intestine to the stomach
in the faeces. Between egg and adult are a number of and this induces ripe eggs to hatch, releasing
larval stages and these may occur in vertebrates or infective hexacanth larvae that give rise to cysticerci.
invertebrates according to the worm species. Reports emerge from time to time about the efficacy
of a tapeworm infection as a slimming agent, which
Pork (Taenia solium) and Beef (Taenia saginata) it might be, but as you can see that is only half the
Tapeworms story!
These are so similar that they may be described
together. Taenia solium is about 4 metres long and Phylum Nemertea or
Taenia saginata is about 6 metres long. Both have Rhynchocoela (Proboscis or
two hosts in the life cycle and both hosts are ribbon worms)
vertebrates. The host to the adult worm is always
human. Symptoms of infection include diarrhoea
and weight loss. The adult worm releases eggs that
 27

Rhynchocoela means “beak” or “snout cavity”, in system, which is an advance over flatworms. There
reference to this worm’s proboscis: Ribbon worms is however, no heart and blood flow is not strictly
have a proboscis or snout that is eversible so that it controlled. Contractions of the body musculature
can be projected forward to catch prey. Some ribbon play a large part in moving blood around the
worms have a puncturing stylet (“arrow”) at the tip vessels.
of the proboscis to inject poison into their prey. This
The organisms (worms) discussed in this
proboscis is usually stored inverted in the
chapter represent an increase in complexity over
rhynchocoel. Some 1 300 species of ribbon worms
the Cnidaria; for instance, they all have excretory
have been described. Most are free living, but a few
systems. Advances are most apparent in the ribbon
have taken up a parasitic or commensal lifestyle.
worms whose elaboration of a blood system and
Ribbon worms vary in length from a few one-way intestine is also exhibited by higher
millimetres to some 30 metres; the longest ribbon animals. The solid triploblastic body plan remains
worm, Lineus longissimus, is probably the longest a limitation, however. Movement of the circulatory
animal on the plant. They are variously coloured fluids and the gut contents are largely determined
from inconspicuous pale hues to mixed patterns by whole body movements and, to a lesser extent,
with orange, red, green and yellow predominating. the body is influenced by movements of the muscles
Most are marine, some live in fresh water and a few surrounding the gut and circulatory system. Higher
are terrestrial. efficiency might be gained if the body layout
allowed more independent movement of the organs
Although nemerteans have no special economic
and systems. The next group, the nematodes or
or medical importance, they are biologically
roundworms, achieve this with a primitive cavity
noteworthy for their anatomical features that place
called a pseudocoelom.
them in an ascending order of complexity from
flatworms to higher animals. They possess a
oneway gut, which allows its different parts to be
optimised for different digestive tasks: this is in
contrast with the blind-ended gut of flatworms. The
acoelomate layout in common with flatworms
means that movement of material in the gut is still
achieved largely by the contractions of the entire
body rather than by specialised gut musculature.
Also in common with turbellarians, they have a
ciliated ectoderm, and some ribbon worms can
reproduce asexually by fragmentation as well as
sexually with gametes. In contrast with flatworms,
ribbon worm sexes are separate (dioecious).
Furthermore, they have a primitive circulatory
 28

as a pseudocoel or pseudocoelom. A pseudocoel is a

cavity lined by mesoderm (and thus muscle) on its
outer side only. Because the pseudocoel is filled
TRIPLOBLASTIC WITH A FALSE BODY CAVITY
with fluid, the cavity can act as a hydrostatic
skeleton. Unlike the other “worm” phyla, nematodes
ECDYSOZOA (ANIMALS THAT MOULT): belong to the group Ecdysozoa as opposed to

Phylum Nematoda Lophotrochozoa, because of the tough inelastic

cuticle, moulting (ecdysis) takes place between
The nematodes - roundworms and
larval stages, of which there are normally four.
threadworms - are all long, thin, spindle-shaped
worms. Although they all look rather alike, the tens Most pseudocoelomates have both circular and

of thousands of species of nematodes are adapted to longitudinal muscles in their body walls but

a wide variety of biotopes. They are found virtually nematodes have a rather different system. While

everywhere, from jars of vinegar to beer malts, from they have well developed longitudinal muscles, they

soil and litter to the deep ocean, from hot springs at have no circular muscles at all. The antagonist of the

60oC to the freezing Antarctic, and from the longitudinal muscles is no more than the

haemocoels of insects to the roots of plants. Indeed, pseudocoelomic fluid, which is kept under high

it has been said that if one were to remove pressure (up to 0.5 atmospheres or 225 mm Hg,

everything else on Earth and leave only the depending on species) by a series of inelastic fibres

nematodes, one would be able to discern a vague wound spirally around the worm to form a tough

outline of the Earth and most of its inhabitants. cuticle. This cuticle prevents the worm from
changing length so that when the longitudinal
Most nematodes are extremely small (<<1 mm
muscles contract on one side, the worm bends to that
long) so that, although they need to live in water,
side. When the same muscles relax, the high pressure
they can comfortably survive in the thin film around
of the body fluid returns the body to its original
individual sand grains in soil, where densities may
shape. As a result of this rather strange arrangement,
be as great as 105 in a spoonful of soil. Nematodes
nematodes are able to move, although they are
living at the bottom of the sea or of lakes, and many
restricted to a series of sinusoidal or snakelike
parasites, may be larger (up to about 100 mm long),
movements, which they make by contracting the
while the giants of the nematode world are those,
muscles alternately on one side of the body and then
like Ascaris, that parasitise the guts of vertebrates.
the other. They are very effective at moving between
Taxonomically the nematodes are divided into sand grains, and some can writhe so rapidly that they
two classes and numerous families, genera and can effectively swim, but none is by any means an
species, but the taxa are not easy to tell apart without athlete of the animal kingdom. (Their simple form
the help of genetic tools, and we will not consider and limited locomotor abilities are ideal for a
their taxonomy any further. parasitic existence, though.)
Nematodes are triploblastic animals with a
mouth and anus, and a fluid-filled body cavity known
 29

Some aspects of nematode architecture are truly reproductive system hold the vulva of the female
bizarre. Firstly, the longitudinal muscle layer is only open and the sperm are introduced into the uteri
one cell thick, but the muscle cells are large and under pressure. Eggs are usually protected by a hard,
complex. Secondly, instead of having a series of resistant shell and released into the environment,
nerves serving these muscles, each muscle cell sends where they may lie dormant before hatching to form
out a long process that attaches to either the dorsal larvae that look like miniature adults. In some cases,
or the ventral nerve cord, which run down the length the eggs are covered by desiccation-resistant cysts
of the body. Cephalisation is minimal and, although and the young larvae inside can survive long periods
there is an anterior nerve ring, nematodes do not of desiccation and extremes of temperature.
have a true brain.
Another unusual feature of the nematodes and
The high internal pressure has all sorts of odd their allies is the phenomenon of eutely: the
consequences. Just think of how difficult it must be number of cells in each organ is fixed within a
to take food in, or to prevent food in the gut (or eggs species. At each larval moult the number of cells in
or sperm) from being shot out of the body by the high certain organs increases until after the fourth moult
pressure (but how easy it must be to defecate). The the invariable adult number is reached. The entire
combination of high internal pressure and a lack of excretory system of most nematodes consists of only
jaws restricts most nematodes to feeding on liquids one or two cells.
or very small particles of organic debris or living
Many, many species of nematodes are parasitic on
algae (a few are predatory, though). The pharynx is
plants or animals. Generally, their life cycles are
muscular and pumps food into the oesophagus.
simple and most have a single host. Ascaris
Because the gut lacks muscles, food moves through
lumbricoides (the human and pig roundworm) is a
it merely by the pumping action of the pharynx. The
major intestinal parasite of humans and is the largest
anus and the genital opening are kept closed by
nematode known. Eggs are excreted in the faeces of
particularly effective sphincter muscles, and
the host and are extremely resistant to damage.
defecation is explosive.
(They can even survive storage in formalin.)
Most nematodes are dioecious, although some are Infection occurs when humans or pigs eat food
hermaphrodite and a few, which live in alternately contaminated with faeces and thus with eggs. Ascaris
wet and dry habitats, alternate a parthenogenic is generally not dangerous to its host unless the host
(developing from an ovum without fertilization) and is poorly nourished, or the parasite load is extremely
a sexually reproducing stage in their life cycles. The high (in which case the parasites may block the gut
male reproductive system consists essentially of a and so kill the host). The overall effect on human
testis and a vas deferens. Nematode sperm lack populations can be devastating, though. It has been
flagella and are probably unable to move by estimated that in China, for example, more than 10%
themselves. The female reproductive system of food consumed goes to nourishing human
consists of paired ovaries, oviducts and uteri. parasites, mostly Ascaris. Other debilitating and
Copulatory spicules at the tip of the male's sometimes lethal parasites of humans include
 30

pinworms, hookworms and Dracunculus, as well as sipunculids. Sipunculids (peanut worms) were
filarial parasites that cause river blindness and previously classified as a separate phylum but is now
elephantiasis. included under Annelida, however, this group will
not be discussed in this course. Leeches hold a
peculiar fascination because of the habit of some of
Minor pseudocoelomate phyla them to feed on human blood. However, in global
The pseudocoelomate condition also occurs in a terms the oligochaetes (earthworms and their allies)
number of smaller phyla, consisting of few species, play a far more significant role in turning over and
that are typically small and cryptic, and mostly form aerating the soil, while polychaetes are often the
part of meiofaunal communities. They include the most abundant organisms in soft marine sediments.
minute rotifers, gastrotrichs, loriciferans and It has been estimated that there are 22 000 species
kinorhynchs (all mostly in fresh waters), the squat, of annelid worldwide, with about half of these being
wormlike marine priapulids, and the elongate polychaetes.
wireworms of the freshwater phylum Annelids are metamerically segmented
Nematomorpha. Of all these groups, the Rotifera are triploblastic coelomates with a complete gut (i.e. a
the most common and ecologically the most mouth and anus), a closed blood system and a
important. They live mostly in fresh waters, although ganglionated double ventral nerve cord.
some are found in the sea. They feed on Metameric segmentation is the state of having the
phytoplankton using a distinctive ciliated wheel body divided into a series of similar units, or
organ, which is also used for locomotion. Some are segments, each containing a part of the coelom. Thus
planktonic but many of them spend most of the time these animals consist of a head followed by a series
attached to the substratum by means of a pair of of units repeated down the length of the body. The
“toes”. coelom is filled with fluid and functions as a
hydrostatic skeleton, just as it does in many other
groups of animals. In oligochaetes, and most
polychaetes, each segment is separated from the
THREE LAYERS AND A BETTER BODY CAVITY: next by a transverse septum stretching right across
TRIPLOBLASTIC COELOMATE ANIMALS the body. This means that each segment is a
selfcontained unit with a separate hydrostatic

LOPHOTROCHOZOA: skeleton. Since each segment is also provided with

a nerve ganglion, it can act more or less
Phylum Annelida independently of the others. This allows peristalsis
to occur and so most annelids can ooze along on the
The phylum Annelida includes the errant and
substratum, and many can burrow into soft
sedentary polychaetes (class Polychaeta), the
sediments. The bundles of hard, chitinous setae
clitellate worms namely oligochaetes and
found ventro-laterally and dorso-laterally on each
hirudineans or leeches (class Clitellata), and
segment of oligochaetes and (errant and sedentary)
 31

polychaetes assist in these movements by acting as circumoesophageal commissures to a

small anchors. suboesophageal ganglion, which controls much of
the feeding activity. A ganglionated double ventral
Because annelids are coelomates, the gut is
nerve cord sends lateral nerves to each segment.
muscular and innervated. This means that it can pass
Some of the largest giant fibres known (up to 1.5 mm
food down its length from mouth to anus. It can also
in diameter, found in the polychaete Myxicola
develop separate sections, like a muscular gizzard
infundibulum) provide a rapid escape (retract)
for grinding food, and a crop for storing it. Most
response for tube-dwelling polychaetes. Some
annelids have salivary glands that produce an
polychaetes have eyes or simple light receptors, but
enzyme-containing lubricant, but none has the
it seems that annelids do not respond to sound. We
equivalent of a liver. Instead, the gut wall looks
know this because of the work of Charles Darwin,
yellowish-brown because it is covered by
who wrote his last major work on earthworms. He
chlorogonenous tissue, which acts as a producer
noted that he had played a piano and a bassoon to his
of enzymes and also a storage organ for both food
earthworms, and whistled and shouted at them, but
and harmful particles. Many polychaetes have an
to no effect.
anterior proboscis that can be extended in front of
the head and used either for feeding or as an anchor
for burrowing.

Annelids other than leeches have two separate

fluid-filled systems within the body. The one is a
closed blood vascular system, which consists of a
dorsal contractile vessel and a number of peripheral
blood vessels that end in capillary beds. The blood
itself often carries a respiratory pigment (red
haemoglobin, green chlorocruorin or
reddishbrown haemerythrin). Parapodia,
tentacles and other extensions of the body are filled
with blood and act as gills. The other fluid-
containing system is the coelom which drains into
nephridiopores and opens to the outside (of the
body) by means of a pair of coelomoducts (or
nephridia) in each segment. Fluid balance is
maintained by means of a pair of nephridia in most
segments.

The nervous system of most annelids is fairly

sophisticated. A cerebral ganglion (a “brain”) is
linked around the oesophagus by two
 32

when the right-hand circulars contract, the left-hand

end bulges. In D, the right-hand circulars contract
Hydrostatic skeletons and all the others are relaxed; pressure is dispersed
equally throughout the body, which becomes both
Worms and many other animals have no hard longer and thinner.
skeletons or limbs that would allow them to swim, or
crawl. Instead, they rely on a very simple physical POINTS TO NOTE:

principle: water, like other fluids, is incompressible.

1. Muscles can do work by exerting force when
Water can therefore be used as a “hydrostatic
they contract but NOT when they relax.
skeleton”. Imagine a sausage skin made up of
longitudinal and circular "muscles” and filled with 2. Muscles cannot lengthen themselves. The
water. If the longitudinal muscles contract while the only way that a muscle can regain its resting length
circular ones relax, the pressure is transmitted is by being extended by an opposing force. This is
throughout the incompressible water and the usually another muscle. The biceps and triceps
sausage will become short and fat. If the circular muscles in your arm act as antagonists to each other,
muscles then contract while the longitudinal ones as do the circular and longitudinal muscles in the
are relaxed, the pressure of the muscles will be body wall of a worm.
transmitted throughout the water and will cause the
whole sausage to become long and thin. That is
exactly how hydrostatic skeletons work: a simple,
worm-shaped body can change length and can thus
crawl or even burrow. A hydrostatic skeleton
principle is also used for different activities and
structures in other animals too. The tube feet of an
echinoderm, the proboscis of a polychaete or a snail,
the foot of a bivalve, the legs of millipedes and
onychophorans, and even the expulsion of
mammalian foetuses from the uterus, rely on
hydrostatic skeletons of one sort or another.

The cylinders below (Fig. 7) represent a worm

Fig. Schematic diagram describing the
with longitudinal and circular muscles in its body
antagonistic working of circular and longitudinal
wall. In A, both are relaxed. In B, the longitudinal
muscles to allow movement.
muscles are relaxed while the circular muscles to the
left maintain tension and the circular muscles to the
right contract, so the right-hand end elongates. In C,
all the longitudinal muscles maintain tension so that
 33

Class Polychaeta (Bristle worms) worm spawns on the ¾ of full moon in July; Pacific
islanders harvest these epitokes (egg cases) as a
The class Polychaeta is divided into a number of
delicacy. A fertilized egg develops into a topshaped
subgroups: we distinguish (informally) the Errantia
trochophore larva, which is planktonic and
(“wandering”) with a number of subclasses; the
planktotrophic (i.e. lives and feeds in the plankton),
subclass Sedentaria (“stationary”); and the subclass
before settling on the bottom to grow to adulthood.
Echiura, or spoon worms (the latter will not be
Molluscs also have as their earliest larva a
discussed further in this course). The classification
trochophore, which implies a relatively close
of Errantia is currently being debated, but for the
evolutionary relationship between the two phyla. (It
purposes of this course we will treat this group as a
is possible, though, that the similarity is functional
subclass.
rather than evolutionary.)
“Subclass” Errantia (Errant Polychaetes)

The errant polychaetes, or bristle worms, are the

Subclass Sedentaria
commonest worms in marine environments.
Internal anatomy is generally similar throughout the (Sedentary Polychaetes)

group and they all have parapodia (lateral flaps) The sedentary polychaetes usually live in
bearing numerous setae. Using their parapodia and burrows or tubes, which may be reinforced by
the muscles of the body wall, some errant proteinaceous or calcareous bodily secretions or by
polychaetes can even swim by throwing their bodies small sand grains. They are particle feeders, either
into snake-like curves and using the parapodia as filtering particles from the water or collecting
paddles. The errant forms, because they move deposits from the surface of the sediment, usually
around, generally have a distinct head with tentacles, using ciliated tentacles or mucus to do so. They have
eyes and palps, with well-developed parapodia used no obvious head, either because it is indeed tiny, or
for crawling and swimming. They tend to be grazers because it is obscured by the presence of large and
or detritivores like Pseudonereis variegata, or complex tentacles and gills. In general, the parapodia
predators like Eunice aphroditois, which is known to are reduced, although the bundles of setae may be
anglers as “wonderworm” and can grow to a length well developed and assist the animal in moving up
of a meter or more (and can give the unwary handler and down the tube or burrow. Well known South
a nasty bite). African forms include the bloodworm, Arenicola

Except for some rare freshwater forms, errant loveni, which burrows into sand flats at low tide, and

polychaetes are all dioecious. They have no distinct Gunnarea capensis, which makes huge reefs of sandy

reproductive systems, eggs or sperm being produced tubes at high tide on the west and south coast.

in the lining of the body wall. When it is time to lay

eggs, in some species, all members of a population Class Clitellata: Oligochaeta and
simultaneously swarm towards the surface of the Hirudinea
sea, where they release their eggs or sperm in a The oligochaetes and leeches have far more in
process known as broadcast spawning. Palolo common with each other than either does with the
 34

polychaetes. Both are more commonly found in fresh found near sewage outlets and other sources of
water or in soil than in the sea; members of both lack organic pollution. One, a small (~ 10 mm) worm
parapodia but develop a clitellum, at least during the called Tubifex, is bright red because it contains
breeding season; both oligochaetes and leeches are haemoglobin. It occurs in countless millions in
hermaphrodite, have complex reproductive anaerobic muds.
systems, copulate to exchange sperm, and protect
their eggs in cocoons; none has a trochophore larva.
These similarities suggest that the two subclasses Subclass Hirudinea (leeches)
are closely related. Indeed, it seems that leeches Anatomically the leeches are very odd annelids,
evolved many millions of years ago from a group of the coelom being reduced to a series of sinuses and
ectoparasitic freshwater oligochaetes. the body filled with a tough, fibrous tissue known as
botryoidal tissue. The explanation seems to be that,
while most present-day leeches feed on a variety of
Subclass Oligochaeta
small invertebrates, the group originated from some
The oligochaetes are the earthworms and their
oligochaete adapting to an ectoparasitic way of life.
allies. They are typically wormlike, lacking an
A parasitic leech like Hirudo medicinalis spends
obvious head or parapodia and bearing only a few
much of its time attached to the substratum by
setae in four bundles per segment. All oligochaetes
means of its posterior sucker. When in need of a
have complex reproductive systems and most can
meal, it stiffens its body and stretches out into the
also undergo asexual reproduction by splitting or
water, waiting to touch and latch on to a potential
budding. Although some occur in fresh water and
host. It attaches by both anterior and posterior
others in the sea, these aquatic forms are generally
suckers, makes a small incision in the skin of its host,
small and cryptic, living burrowed in sediments. In
secretes an anticoagulant substance called hirudin
contrast, terrestrial earthworms such as Lumbricus
(and a local anaesthetic so that the host does not feel
terrestis are of major importance in turning over and
pain), and takes in a massive blood meal. When it is
thus aerating soil.
full, it drops off the host and crawls away to a safe
In undeveloped parts of the country, where
retreat, where it will spend several months digesting
summers are very dry as well as hot, earthworms are
its meal. Leeches feed on any vertebrate host,
not particularly common. Many of the species of
including fish and sea turtles, although H. medicinalis
earthworm now found in cultivated and irrigated
prefers terrestrial mammals. This medicinal leech
soil have been accidentally introduced to the region
can take in up to ten times its body weight in blood
from Europe. An impressive native species is
and can survive on a single meal for up to a year. It
Microchaeta, a giant earthworm that grows to over a
secretes no digestive enzymes whatsoever, and for
metre in length and the thickness of one's thumb, is
digestion of its food relies entirely on symbiotic
found after rains in the Eastern Cape around
bacteria in the gut. For several centuries, the ability
Grahamstown. Many aquatic oligochaetes are fairly
of leeches to take in such huge meals was exploited
resistant to pollutants, often being the only animals
 35

by doctors practising western medicine, who SOFT-BODIED, TRIPLOBLASTIC COELOMATE

believed that many illnesses were caused by an ANIMALS, WITH SHELLS
excess of blood. Interestingly, medical doctors
presently use leeches to drain very badly bruised
Phylum Mollusca
tissues, as well as extracting hirudin for its
This phylum is the second largest, after the
anticoagulatory properties.
arthropods, housing about 100 000 species. Some
All leeches, whether ectoparastic or not, move by
are of great significance to mankind, as pests, disease
means of a looping motion. A leech will first attach
carriers or a source of food. Body form is diverse,
its posterior sucker to the substratum then will
including creatures as different as snails, mussels
stretch its body as far forwards as possible before
and octopi. This diversity makes it very difficult to
attaching its anterior sucker to the substratum. It
produce a single definition that includes all forms. All
then releases the posterior sucker and shortens its
are triploblastic (with three body layers); the body
body, bringing the posterior sucker close up behind
is soft and unsegmented and has a through-gut
the anterior one and attaching it once more.
with a mouth and anus. Most species are bilaterally
The anatomical peculiarities of the leeches make symmetrical (obscured by a “twisting” of the body
perfect sense in the light of these adaptations to in one group - gastropods). The majority have an
feeding and locomotion. The gut has to be massively external shell made of calcium carbonate, which is
expansible because of the huge blood meals it can produced by the epidermis. The body usually has
accommodate; locomotion is by looping but most three regions: a foot for locomotion, a head with the
often when the animal is active at all, it is stretched mouth and sensory organs, and a visceral hump
out waiting for prey. In either case, the body acts as that houses the digestive, excretory and
a single unit, so a large coelomic space and reproductive organs. The body is covered by a
independent segments would result in an inefficient mantle that forms a cavity sheltering a pair of
use of energy. And, of course, parapodia and setae chemosensory organs called osphradia and a pair of
are not needed for this way of life. respiratory gills (which have a unique ciliated

Although large, blood-sucking leeches are most structure and are called ctenidia to distinguish them

common in the tropics, old records do exist of from other gills). At the front of the gut is a long

medicinal leeches in rivers of the southwestern ribbon-like structure with rows of teeth, termed the

Cape. Much smaller leeches, particularly predatory radula. The coelom is usually reduced to small

forms, are common in South African fresh water cavities around the heart and gonads. The blood

systems. A few species are also known from inshore system is typically open: the heart pumps blood into

marine environments. arteries that open into a diffuse haemocoel, which is

the main body cavity. The nervous system consists of
a ring around the oesophagus and nerve cords
running down each side of the body. Usually the
 36

nerve cells are concentrated into ganglia, plates into which the shell is divided. Chitons are
particularly in the head. primitive, bilaterally symmetrical, and have a
mouth at the front end, a long, coiled gut and a
In the more primitive molluscs, fertilisation is
posterior anus. A large flat foot used to attach firmly
external and leads to a top-shaped trochophore
to rocks. Almost all are grazers, using a powerful
larva that is planktonic and has two rows of cilia
radula to scrape algae from the substrate. Chitons
that propel it through the water. This closely
are considered primitive, resembling the ancestral
resembles the larva of annelids (and is one reason
mollusc form.
the two phyla are suspected of having a common
ancestor). The trochophore turns into a veliger
larva, which is also planktonic, and has two large
Class Bivalvia
anterior ciliated lobes (called the ‘velum’), and a Most mollusc classes have highly modified body
small shell. Eventually this larva settles on the sea shapes. However, clams, mussels and oysters, who
floor and metamorphoses into a miniature version of all belong to the class Bivalvia, are characterised by
the adult. having two valves constituting the shell, joined
together middorsally by a springy ligament and
There are eight classes in the phylum. Two are
positioned laterally on either side of the body. The
rare and are not dealt with here. The class
fact that the valves are lateral (so a left and right
Monoplacophora is also unlikely to be encountered,
shell) is important, because it distinguishes the
but of such interest that it deserves mention.
Bivalvia from an entirely different but similar
Monoplacophorans have a helmet-like shell, and
looking phylum, the lamp shells or Brachiopoda,
their internal organs are repeated in a manner that
which have dorsal and ventral valves covering the
once raised the (incorrect) idea that they were the
body. Bivalves are completely encased in their shell
only segmented molluscs. The discovery of living
and are relatively sedentary. Clams live in sand or
specimens in the deep-sea in 1952 created great
mud and have a large wedge-shaped foot that allows
excitement because they were previously known
them to burrow. Scallops are considered free-living
only as fossils and thought to have gone extinct in the
being able to move short distances by flapping the
Devonian period. Modern species are thus living
shells. In contrast, mussels produce a beard-like
fossils. Another class that can be mentioned briefly is
byssus for attachment to rocks, whereas oysters
the Scaphopoda, all of which have tusk-shaped
cement one of their shell valves to the rock face. In
shells, a short foot for locomotion, and a series of
all bivalves, the head is reduced and does not have
club-tipped tentacles on the head for feeding. The
eyes or sensory tentacles.
remaining four classes demand more attention.
Nearly all bivalves have very large ctenidia that lie
Class Polyplacophora in the mantle cavity, used for both respiration and
filter-feeding. This method of feeding is probably the
Sometimes called Amphineura, this class contains
reason why no bivalves live on land. Cilia on the gills
the chitons. The name Polyplacophora literally
pump water into the mantle cavity via an inhalant
means “many-shell bearer” and refers to the eight
 37

aperture, and over the gills. There, the cilia capture Class Gastropoda
tiny particles from the water and drive them towards
This is a very large group of about 40 000 species
the mouth. Labial palps on either side of the mouth
and includes the snails, winkles, whelks, limpets
sort the particles and transmit the edible fraction
and sea slugs. Most are marine, but representatives
into the gut. Waste water and unwanted particles are
can be found on land and in fresh water; this is the
expelled through an exhalant siphon at the back of
only molluscan class that occurs in all three
the body. Inside the stomach is a long thin rod of
environments.
concentrated enzymes (called the crystalline style),
and this is constantly rotated to grind away the tip, The gastropod shell is a single structure and is

releasing enzymes to digest the food. Bivalves are typically coiled in a spiral. The reasons for this are

the only molluscs to lack a radula. deep-seated, however, it makes the shell more
compact. If it gets too big, it leans over to one side
Most bivalves practise external fertilisation and
(generally the right) to balance the weight of the
go through trochophore and veliger larval stages,
coiled shell, producing a shell with a pointed tip
but a few retain their fertilised eggs inside their
rather than the shell of a nautilus (the latter is not a
mantle cavities and brood them there until
gastropod).
miniature adults are released.
A more peculiar process specific to gastropods is
Oysters are an important commercial product,
torsion. During larval development, torsion takes
and they are cultured world-wide for food and some
place, and powerfully affects the body structure of
to produce pearls. Mussels, scallops and clams are
adults. Torsion involves twisting the body so that
also seafood delicacies.
the top half (the visceral mass) rotates through 180°
Mussels have the capacity to dominate wave- relative to the foot and head, thus swinging the
swept rocky shores and are often the most mantle cavity and anus from their originally
abundant creatures there, often providing a posterior position and leaving them positioned over
substrate for other species like polychaete worms the head. One theory is that this allows the veliger
to crawl in between the byssus threads. An alien larva to pull its body more easily back into the shell,
species, the Mediterranean or blue mussel (Mytilus thus decreasing the chances of vital and soft body
galloprovincialis), has been introduced to South parts being snapped-at by passing predators.
Africa in the last few decades, and is aggressively Another theory is that torsion is beneficial because it
invading the coast. It now dominates most wave- moves the mantle cavity, and gills with its
beaten rocky shores on the West Coast and is chemosensory osphradia up front. Torsion,
rapidly spreading, having displaced the native however, also creates problems: now the anus is on
Perna perna (Brown mussel). Mytilus top of the head end and there is a risk of fouling the
galloprovincialis is preferred over the local species gills and other sensory organs.
by the mariculture industry that supplies local
Gastropods have evolved all sorts of tricks to
restaurants, because it is plumper and faster
overcome this problem, but four of these are typical.
growing.
1) The holes that run down the one side of a
 38

perlemoen (abalone) shell are exhalant openings Subclass Prosobranchia (Whelks, winkles
and limpets)
that deflect faeces and wastewater material out to
one side to avoid contaminating clean incoming The most typical and familiar gastropods belong
water that oxygenates the gills. Other gastropods in this subclass. They are marine and retain torsion
have solved the problem differently. 2) Keyhole and a twisted, asymmetrical nervous system. The
limpets evolved directional flow over the gills, mantle cavity opens above the head (the name
through the mantle, past the anus and out through a Prosobranchia translates to “front gills”). In the
“chimney” on the top of the shell. Furthermore, 3) primitive condition, the mantle contains two gills,
many gastropods, such as whelks, have lost the one although one is lost in the more advanced forms. The
(right) gill to allow for the incoming current from the sexes are usually separate. Fertilisation is external in
siphon to run over the left (single-branched) gill primitive species, which undergo a full larval
first, then past the anus and out the exhalant side. development with planktonic trochophore and
Finally, 4) the sea hares detorted either 90o or 180o veliger stages. Many of the advanced forms have
with the anus on the side or back end again. internal fertilisation, and the larval stages are often

Most gastropods have a large foot for locomotion. reduced, with either the trochophore or the entire

The posterior surface of the foot carries an larval development-taking place inside an egg case.

operculum that acts like a door when the animal The habits of individual species vary greatly, but
pulls back into its shell, blocking the entrance to the in general, the more primitive forms such as the
shell and protecting the animal from predators or winkles and limpets are herbivorous, using the
desiccation. The head is well developed, containing radula to feed on seaweeds or to scrape the rocks to
eyes and sensory tentacles. gather microalgae and sporelings of seaweeds. The

The blood system is well developed and there is herbivorous nature of these species is easily

usually a three-chambered heart with two detected, for they all have round apertures to their

auricles (receiving blood) and a ventricle. The shells. Limpets exert a powerful influence on the

excretory function of the kidney is linked to the nature of rocky-shore intertidal ecosystems,

beating of the heart. When the heart contracts, the controlling seaweeds. South Africa’s limpets are

pressure within it cause blood plasma to be filtered more diverse and more specialised than anywhere

through the wall of the heart into the pericardial else in the world. Some enjoy a specialised

cavity, from where it passes through the relationship with particular seaweeds, and create

renopericardial canal and into the kidney. There, tiny ‘gardens’ of these, which they defend against

useful substances are reabsorbed, and more wastes other grazers, fertilise and weed to prevent

are added from the blood. The final urine is excreted overgrowth by other larger seaweeds.

through the nephropore into the mantle cavity. The more advanced prosobranchs include the
whelks, and most are carnivores. The possess a
long siphon that projects forwards out of the mantle,
and draws in water to their osphridia, to sense prey
 39

ahead. Because of this, the aperture of the shell is nematocysts from their cnidarian prey and use them
always elongate and notched at the front to house as a second-hand defence against would-be
the siphon. This gives a convenient clue to the predators. To advertise their “nastiness”, they have
predatory habits of the animals. Several whelks drill vivid colours. Species in one well-known group, the
neat holes through the shells of their prey, using a Nudibranchia, are popular with divers and
combination of their radulae and chemicals that aquarists because of their gorgeous colours and
dissolve the shell. One species, Argobuccinum, elaborate body shapes.
produces sulphuric acid (pH 2.0!) to dissolve the
The “Pulmonata”
sandy cases in which live their main prey, Cape reef
This group includes the land and freshwater
worms.
snails and slugs. Members of this subclass are
Subclass Heterobranchia (different-gilled nearly all simultaneous hermaphrodites (having
snails)
both male and female organs at the same time), have
The “Opisthobranchia” (sea snails and fragile or reduced shells, and have lost their original
nudibranchs)
respiratory organs, the ctenidia. There are several
These animals have reduced or lost their shells superorders within the subclass, but two are worth
and lost their ctenidia. They have also undergone singling out.
“detorsion”, a reversal of the process of torsion - so
They have all lost their ctenidia and converted
that their mantle cavities are again posterior or at
their mantle cavities into airbreathing “lungs”,
least lie on the side of the body (the name
richly lined with blood vessels to absorb oxygen
Opisthobranchia means “posterior gills”). The body from the air. Land snails retained their spiral shells,
becomes almost bilaterally symmetrical as a result, although they are very fragile, probably reflecting
losing its coiling. In the process of these evolutionary the difficulty of getting enough calcium to
steps, some of the opisthobranchs that have lost manufacture strong shells away from marine
their ctenidia, re-evolved a new and different set of systems. Land slugs produce no shell at all. The
gills coming of the back of the anus. All retain a doorlike operculum has also been lost in
twisted nervous system as a memento of their torted pulmonates. A single genus, the limpet-like
ancestry. Siphonaria, has returned to the sea and has both
Why should animals evolve a pattern like torsion lungs and secondary gills, so it can respire in water
and then give it up to become ‘detorted’ once again? and air. Not surprisingly, hermaphroditic land and
We really don’t know, but one theory is that the freshwater pulmonates sperm-swap, but have no
ancestors of opisthobranchs became burrowers, and free swimming larvae. Larval development takes
a shell and anterior mantle cavity would have been a place in large yolky eggs (lecithotrophic eggs) that
distinct disadvantage while ploughing through the hatch into miniature adults.
sand. Whatever the explanation, the reduction or Most pulmonates are grazers, and several have
loss of a shell leaves their soft bodies exposed, and become serious agricultural pests after being
many have developed toxic chemicals or ingest introduced to foreign lands. The garden snail, Helix
 40

aspersa, which is so common here now, was octopus), has a pretty, paper-thin external shell,
originally introduced in the mistaken belief that it which is avidly hunted by beachcombers in summer
was the edible snail so popular in Europe (a larger when storms wash them ashore. This shell is in
species in the same genus). It has now spread and reality the egg case of the female and bears no
become a notorious garden pest. Other pulmonates resemblance to the true shells of other molluscs. It
are important carriers of disease, the most notable seems as if the paper nautilus has gone through an
being the freshwater snail, Bulinus, which transmits ancestral stage like the octopus, completely losing
bilharzia. the ability to produce a molluscan shell, and has then
evolved a secondary shell that is a completely new
Class Cephalopoda (inkfish) invention. The male paper nautilus is minute, shell-

This class includes the squid, cuttlefish and less, and is often found inside the shell of the female.

octopus, which are among the most advanced and In all cephalopods, the mantle cavity is enlarged
largest of all invertebrates. Squid reach up to 20 m in and the walls strongly muscular, drawing in water
length! The body is modified from the basic and pumping it over the gills that are housed inside.
molluscan plan, with parts of the head and foot This water can be powerfully expelled through a
fusing to form 8 or 10 long tentacles armed with siphon, achieving locomotion by jet propulsion.
rows of suckers. An external shell is retained in only Cephalopods are very active animals with a high
a single group, represented nowadays by the pearly oxygen demand. Their blood systems are complex,
Nautilus. Two of the three subclasses, Nautiloidea with two separate sets of hearts, and the vascular
and Ammonoidea, were once dominant groups and system is closed (unlike all other molluscs), making
there are abundant fossils testifying to their it a much more efficient means of transporting
previous success. Both these subphyla had large, oxygen.
flatly coiled external shells, and some were relative
Cephalopods are all predators, capturing fish,
giants, with shells over a metre in diameter.
crustaceans and other molluscs with their tentacles
However, the ammonites went extinct at about the
and shredding t hem with their beaks and radulae.
time dinosaurs were disappearing from the face of
They have extremely well-developed sensory
the earth, and only a single representative of the
organs. Their eyes are astonishingly like those of
nautiloids, the pearly Nautilus, survives today. All
human beings, with a lens, cornea, iris and
other living cephalopods belong to a third subclass,
diaphragm. This is a remarkable example of
the Coleoidea. The Coleoidea have radically reduced
convergent evolution: two unrelated groups of
or lost their shells. In the cuttlefish the shell is
organisms independently evolving the same
reduced to a chalky internal cuttlebone that serves
structure for the same purpose. Their nervous
as a flotation device, controlling buoyancy. Squids
systems are also highly developed, with massive
have only a horny plate-like remnant of the shell, and
concentrations of ganglia in the head, giving the
octopi have lost the shell completely. The paper
appearance of a brain. This impression is not
nautilus Argonauta (which is badly named because
deceptive as the cephalopods are the most intelligent
it is not a nautiloid but actually closely related to the
 41

of the invertebrates and have a greater learning Abundant and often exquisitely beautiful under
ability than any of the lower vertebrates. the microscope, bryozoans are seldom recognised
because they resemble other organisms, including
They have also developed a system of giant nerve
seaweeds, hydroids and coral. They build colonies of
fibres, which allow very rapid conduction of nerve
minute (1 mm long) individuals, known as zooids,
impulses, quite foreign to other molluscs.
which lie side by side and are individually encased in
All cephalopods practice internal fertilisation.
tiny coffin-like skeletons (called zooecia) made of
The sperm is packaged in very complex
chiton or lime. Each individual is triploblastic and
spermatophores and passed to the female on the
coelomate and has a saclike body with a crown of
tips of specialised tentacles that the male possesses.
filter-feeding tentacles (called the lophophore) that
The eggs are large and yolky, and there is no external
extends through an opening in the zooecium. Some
larval stage; juvenile versions of the adult are
individuals in the colony are specialised into
released. The female often defends the eggs and fans
beaklike structures (called ‘avicularia’), which snap
them to oxygenate them. Growth is extremely fast,
at creatures attempting to settle on the surface of the
and even the giant species tend to mature and
colony. The colonies consist of many hundreds or
reproduce in a single year, often to die immediately
thousands of genetically identical individuals and
after (they are thus semelparous, reproducing only
can form thin crusts on rocks or on seaweeds, or
once and then die).
upright bushy structures, or hard twisted plates, or
Cephalopods are fished as a source of food, and an branching coral-like structures.
important industry has sprung up in the vicinity of
Bryozoans are very common in the sea, and there
Port Elizabeth harvesting squid or “chokka”
are a few freshwater species. They can be found on
(Loligo). They are attracted to lights strung from
the lower surfaces of almost all rocks on the
boats and caught by “jigging” with hooked lines.
seashore, and grow prolifically on various seaweeds,
With all their advanced features, the cephalopods particularly kelp.
must be regarded as the most highly developed
Bryozoans are also commercially important for
invertebrates. Notably. these animals squirt ink for
two reasons. They foul ships and intake pipes and
chemical signalling, and they use chemoreception to
are a nuisance. But some species contain chemicals
disguise themselves or to mimic other species, or to
of great medicinal interest, having anti-cancer
signal, the opposite sex.
properties. Extracts from one species are currently
being tested in clinical trials on humans. Another

Phylum Bryozoa (Ectoprocta) species forms a coating on a whelk and protects it

against predators.
To conclude the section on the Lophotrochozoa,
we will look at a colonial look-alike of molluscs, the
Bryozoa. These are tiny triploblastic coelomates
ECDYSOZOA (ANIMALS THAT MOULT):
encased in shells of different material, thus giving
rise to different shapes and forms. TRIPLOBLASTIC COELOMATE ANIMALS WITH
 42

JOINTED LIMBS The advent of the haemocoel and reduction of the

coelom may be seen as having become possible
because of decreased dependence on a hydrostatic
Phylum (Eu-)Arthropoda skeleton. Such a fluid skeleton was essential to
The (Eu)Arthropoda (“true” arthropods) look movement in the annelids, but arthropods have a
very different from the annelid worms. This is partly hard exoskeleton which allows for a larger body
because they have a hard exoskeleton covering the size and a hard structure to which muscles can be
body and limbs. That part of the exoskeleton attached. A much more efficient use of muscles in
covering the limbs is jointed to allow for movement; locomotion thus becomes possible. The jointed limb
hence the name of the phylum (arthos = joint). The is moved in the desired plane by a muscle anchored
exoskeleton is really a cuticle, the essential firmly within the body and attached to the limb close
component of which is chitin, a to the hinge of the joint, so that the limb magnifies
mucopolysaccharide. However, chitin on its own is the action of the muscle. The limb is thus an effective
soft and pliable; it is made hard by being lever. Complex limb movements are usually
impregnated with calcium carbonate, while in the achieved by using a number of simple joints, each
insects there is incorporation of proteins, which are moving in a different plane. The possession of
then hardened by other substances. These pliable jointed limbs with a hard exoskeleton means that
areas of chitin are the joints and are called movement can be much more elaborate than what is
arthrodial membranes. possible with annelid parapodia. For example, for
Despite differences in appearance and the the first time we meet animals that can use their
presence of a hard exoskeleton, the arthropod plan limbs to raise themselves off the ground, so that
has a great deal in common with that of the annelids. locomotion becomes much more efficient. These
These animals are of course, triploblastic and animals can run, jump and fly instead of just crawling
coelomates and, like annelid worms, they are or creeping.
segmented, they have double ventral nerve cord The exoskeleton, however, does not grow as the
with a ganglion in each segment, and they are animal grows and this implies that an arthropod
bilaterally symmetrical, with a tendency for major must shed its exoskeleton from time to time and
sense organs to be carried on the head. The blood replace it with a larger version. Thus, moulting
system is, however, different. There is a dorsal (ecdysis) is characteristic of arthropods. Further, a
heart with arteries, that open into a body cavity more or less profound metamorphosis may
called a haemocoel; there are no veins and the intervene between the immature and adult stages of
heart simply sucks blood up from the haemocoel some individuals. The sexes are typically separate
through openings known as ostia. As the blood is not (dioecious).
confined to blood vessels, it is an open blood system.
There are several other peculiarities of
The haemocoel is a new cavity not found in annelids. arthropods. One is the compound eye, an eye
The arthropod coelom is reduced to a few small consisting of a large number of units, which are
spaces around the gonads and certain glands.
 43

always in focus. The units are seen on the surface of 3. The segments (tagmata) - instead of
the eye as a series of hexagonal facets. Another repeated like in annelids, are fused and modified to
arthropod characteristic is the total absence of create specialized body regions. For example,
ciliated cells at all stages of development. insects have a head, a thorax and (segmented)
abdomen, whereas many crustaceans have a fused
The arthropods are by far the most successful of
head and thorax to form a cephalothorax. This allows
the invertebrate phyla with > 1 million (M) species
for specialized functions in the fused compartments,
identified and occupying just about every niche, and
mostly containing all the sensitive organs,
a range of habitats; they may well be considered to
specialised appendages, and sensory organs.
outdo the vertebrates themselves and have certainly
been around a lot longer (>150 M years). There are a 4. Specialised respiratory organs – are
number of arthropod sub-phyla. Before we consider present and differ based on the habitat occupied.
the diversity in Arthropods it is worthwhile Spiders have book lungs; insects have trachea and
considering… spiracles and crustaceans have gills. Gills are

Why Arthropods, and particularly insects (on land) efficient in water and spiracles (insects) allow for the
and crustaceans (in the sea) are so successful: direct delivery of air (oxygen) to the tissues via the
tracheal respiratory system. This is particularly
1. The hard exoskeleton - affords fantastic
important for flying insects with a high oxygen
protection for soft body tissue and allows for an
demands while an open blood circulatory system is
increase in body size. The shortcoming of this boxy
slow to facilitate gas exchange.
body design though is that it cannot grow, so need to
moult. During moulting the individual is completely 5. A sophisticated nervous system – that
vulnerable as it cannot move and soft, so an easy covers the entire body through a double ventral
target to predators. The cuticle (especially in nerve cord and branched nerves in each segment.
crustaceans) are not protected with a waxy layer so Specialised sensory organs are connected to this
they may dry out on land. Also, even though the nervous system; the sensory organs include hairs on
exoskeleton allows for a much larger body size than the head and legs to feel vibrations, compound eyes
a hydrostatic skeleton – it is restricted to moderate on the head, with up to 30 000 photoreceptors which
sizes. Very few individuals can grow larger than 1 m provides ~180o vision, or antennae and antennules
in size and they are generally supported by water used for tactile and chemosensory reception.
when this large.
6. Metamorphosis – results in division of
2. The arthrodial membrane – allows the labour with provision and division of resources;
boxy skeleton to articulate across the joints. The this resource partitioning can be between different
large skeleton can articulate, and specialised morphs (body forms) of the same individual or
appendages like wings, legs and antennae can move within a generation with different body forms and
the skeleton efficiently and quickly. It also has complex social structures. In the first instance, the
appendicular specializations like spinnerets to spin same individual will start its life from the egg with
silk, transfer sperm or pincers to bite and grab. resources provided by the mother, to a larva or
 44

young individual which have the explicit purpose to EXTANT ARTHROPODA

feed, grow and then disperse, they may morph into
The extant arthropods can be divided into four
an adult that will reproduce (e.g. egg, to
extant (still living) subphyla. These are: Crustacea,
larvae/caterpillar, finally a moth or butterfly). In
Hexapoda, Myriapoda and Chelicerata. Because
these instances, animals do not compete for the same
the Crustacea and Hexapoda are closely related, they
resources as other size classes as they can time the
are combined in a single clade, the Pancrustacea. The
change-over across these different “generations”. In
Pancrustacea and Myriapoda together form the
the second instance, a single generation yield
Mandibulata. The Chelicerata do not have typical
individuals with different purposes within the same
jaws (mandibles) and are this more distantly related
colony. For example, in the case of bees or ants, some
to the Pancrustacea and Myriapoda.
will become workers, some protectors of the colony
with only one queen to reproduce. In this instance,
there is specialization of tasks and minimising
Subphylum Chelicerata
competition from individuals in the same colony. The chelicerates had their origins in the seas of
the Palaeozoic (about 500 million years ago) but
Given these adaptations it is not surprising that
very few of the present-day species are still marine.
insects and crustaceans are so successful. However,
Only the horseshoe crabs (class Merostomata) and
not all groups have been equally successful over
sea spiders (class Pygnogonida) are the truly
time. We should mention the subphylum Trilobita,
marine representatives of the group. Other
which have been extinct for millions of years. But
representatives of the Chelicerata are all terrestrial:
trilobites once were extremely abundant and
Class Arachnida include the spiders (order Aranea),
dominated shallow seas some 500 million years ago.
scorpions (order Scorpionida), ticks and mites
They dominated for nearly 300 M years but went
(order Acari).
extinct in the Permian mass extinction of the.
Lesser-known members include the harvestmen

Subphylum Trilobita † and pseudoscorpions. The popularity of chelicerates

has been summed up by Starr and Taggart (1992) by
The trilobites had dorsiventrally compressed
stating that “…never have so many been loved by so
bodies, oval outline, and three body regions: head,
few”. Contrary to belief though, only a handful of
thorax and pygidium. However, it is the distinct
these species are dangerous to humans, although
three ‘lobes’ of the central part of the body that
their predaceous and often blood sucking lifestyles
earned them the name of Trilobite. They fossilized
do in fact result in them being of some medical
well because of their hard exoskeleton; on a good
importance to humans. Ticks for example serve as
fossil one can even see clearly the individual facets of
vectors (carriers) of microorganisms that may be
the compound eyes. A famous area for trilobite
present on the blood of their hosts and are
fossils is Ceres, in the Western Cape, or Uitenhage
responsible for several serious human-related
area in the Eastern Cape.
diseases. The majority, however, are major
predators of insects and other invertebrate pests,
 45

and as such are responsible for keeping those this includes on the east coast of N. America
populations in check. The only vegetarian members (Delaware Bay) and Japan, where they scavenge on
of this group are the plant-feeding mites. small marine organisms such as clams. The
compound eyes are complex, consisting of a
The Chelicerata differ from Crustacea and
number of ommatidia. The most bizarre
Insecta by the absence of antennae and
characteristic though is that they have turquoise
mandibles. Instead, the first pair of appendages
(blue) blood so no haemoglobin but have
has been transformed to form chelicerae. These
hemocyanin. The amoebocytes in the blood is used
can be of two kinds: they may bear a small pair of
to stop bleeding in horseshoe crabs but in the
pincers at the end, or alternatively they are sharp,
medical industry to indicate bacterial exposure of
hollow and fang-like. The second pair of legs are
supposed sterile packages like vaccines. This
called the pedipalps, and may end in small pincers,
cannot yet be produced synthetically so the blood of
be tactile, or serve as walking legs. All of the other
horseshoe crabs is “milked”.
body appendages form jointed walking legs. There
has been a specialisation of body segments, and
Class Pycnogonida (sea spiders)
their subsequent fusion into two major functional
These marine chelicerates are very slender-
parts. Segments associated with the head,
bodied and have 4-5 pairs of long limbs. They feed on
mouthparts and locomotion have become fused to
soft-bodied invertebrates such as sessile cnidarians
form the anterior cephalothorax, while segments
(Hydrozoa and Bryozoa) by sucking out their body
associated with digestion etc. and locomotion have
fluids. They are rather small but can be quite
fused to form the posterior abdomen. The original
common in certain marine systems where they live
segmentation does not show up apart from the
on the bottom. Although they are placed in the
abdomen of scorpions, which retained this
Chelicerata, their relationships to the horseshoe
primitive condition. The sexes are always separate
crabs and the rest of the Arachnids is not clear.
(dioecious) in all Chelicerata.

Class Merostomata (horseshoe crabs) Class Arachnida (spiders, ticks and mites,
scorpions, pseudoscorpions, harvestmen)
These animals get their name from the very
This is a very large terrestrial group, which all
characteristic carapace shaped like a horseshoe.
have the following features. There are four pairs of
There are considered living fossils being around for
walking legs, and chelicerae and palps are very
450 M years, but now only five living species of
specialised. They are an old group, with fossils dating
horseshoe crabs are left. The five pairs of walking
back to the Carboniferous (360 million years ago).
legs are similar to the chelicera, and posterior to
They apparently were derived from marine
them are a series of flap-like book gills (modified
Chelicerata called eurypterids and may have been
limbs). The body ends in a spine-like process. They
the first animal invaders of land. The book gills,
are quite large (60 cm long). Limulus is the
which are still found in the horseshoe crabs, have
common genus and are restricted to only a few
become modified to form book lungs, but in others
sandy shores where they are reasonably abundant;
 46

trachea have developed. Book lungs are internal tissue damage, respectively. All spiders live on a
and located in pairs on the segments of the abdomen. liquid diet, since the spider pumps digestive
Each lung is an infolding of the body wall and enzymes into the wound of the prey. The resulting
consists of a number of plates (lamellae) that are semi-digested “soup” is then pumped back into the
held apart by bars, for free air circulation. The gut. The spinning organs are located at the end of the
trachea function in the same way as those of insects, abdomen and produce silk. These so-called
but has evolved independently. The heart is a simple spinnerets are linked to very large silk glands.
tube, but the rest of the blood system is a Spider silk is a scleroprotein, which is emitted in
haemocoel. In most species sperm is packaged into liquid form, and hardened by pulling. The silk is used
a spermatophore, which is inserted into the female for various functions: prey capture (e.g. web
genital opening after a lengthy courtship. Sperm building), safety lines, wrapping up prey, and even
transmission is said to be indirect, as there is no tying down uncooperative females during mating.
copulation. A few of the commoner orders are
Order Scorpiones (scorpions)
discussed below.
These are the oldest known terrestrial
Order Araneae (spiders)
arthropods. The long clearly segmented abdomen
With about 30 000 species this is one of the ends in a sting. The paired eyes are situated in the
largest groups of chelicerates. They differ from other middle of the dorsal carapace. While the chelicerae
chelicerates in the presence of silk producing organs are small and hidden, the pedipalps have become
and by the fact that the pedipalps of the male have greatly enlarged and form a pair of pincers/nippers
been modified for sperm transfer. The latter have the for prey capture and manipulation. An unusual pair
end expanded into a bulb, which the male fills with of chemosensory structures, the pectines are
sperm. During the courtship he inserts these one by located on the ventral surface of the body and
one into the female genital opening after he had samples the substrate as it walks over the substrate.
charged them with sperm. The largest species Some species (including a few from the N. Cape and
(tarantulas, and baboon spiders from South Africa) Namibia) are dangerous to humans and their sting
can reach 9 cm in diameter. The prosoma has may be fatal. The more dangerous scorpions belong
several (often 8) eyes dorsally. In some cases, the to a family that it easily recognised by the
eyes are very well developed and can form an image. combination of very slender pedipalps and a stout
The chelicerae have been modified to form hollow “tail” (abdomen). During the elaborate courtship,
fangs, filled by poison glands. Typically, the venom the male manoeuvres the female over the
is not dangerous to humans, but there are spermatophore, which he has been placed on the
exceptions. In South Africa, black and brown widow ground, until it catches on the female genitalia and is
spiders are very common but rarely lethal, but in the taken up.
drier regions the more dangerous cave spider occurs
Order Acari (ticks and mites)
in caves and rock overhangs. Spider venom may be
This group contains many species that are
neurotoxic or cytotoxic, causing either nerve or
implicated in human disease or are pests of crops.
 47

Mites, although small, are probably the most millipedes is that each trunk segment of millipedes
abundant of all arthropod groups. The body does carries not one, but two pairs of walking legs (hence
not appear to be divided, as the abdomen has the name Diplopoda). The centipedes are
fused with the prosoma. The mouthparts fastmoving carnivores, whereas the millipedes
(pedipalps and chelicerae) are all housed in the move more slowly and are all herbivorous. Both
buccal cone. In some mites the chelicerae may be groups are common throughout southern Africa,
adapted for piercing, and in ticks the nipper part of especially in indigenous forests.
the chelicera has become toothed so that it can
anchor the tick into the body of the host. There are Subphylum Hexapoda
four pairs of legs. While most of the parasitic
The Hexapoda constitutes the classes Insecta and
species are ectoparasites, there are several mites
the Entognatha. The latter is a small class (with few
that live internally in the air passages of
species, including the spring tails or Collembola),
vertebrates, and in the tracheal system of insect
and will not be considered further in this course. The
hosts. Many are free-living but have an ectoparasitic
Insecta is a dominant class in the Animal Kingdom in
stage during which they take a blood meal before
terms of numbers and species diversity. The insects
dropping off the host to moult and then become
specifically are estimated to make up about half of
free-living predators. Ticks are permanent
global species diversity, and on land the proportion
ectoparasites and have a soft skin that can expand
is much greater; they have penetrated virtually
after a blood meal. Normally ticks fall off the host
every possible terrestrial and freshwater habitat, but
just before a moult, and then reattach to a new host.
only the edges of the sea.
They can live for over a year in a non-feeding stage
off the host. Very small ticks (pepper ticks) hatch Insects are mandibulate arthropods whose

from eggs, and immediately find a host for their first bodies are divided into three well-defined regions:

blood meal. head, thorax and abdomen. The head, as in all

arthropods, is formed of six fused segments. The
thorax always has three segments and the abdomen
Subphylum Myriapoda
primitively consists of 11, although in some insects
(previously Uniramia)
this is only apparent in the embryo.
There are two important classes of Myriapoda
The head bears compound eyes and a single pair
which are centipedes (Chilopoda) and millipedes
of antennae. Biting and chewing mouthparts, such as
(Diplopoda). Although they are mandibulate
those of the omnivorous cockroaches, are the
arthropods, their appearance is quite different from
primitive type. They consist of a labrum or upper lip,
that of any crustacean. The body consists of a head
a hypopharynx, paired mandibles or jaws, paired
followed by a long trunk of essentially similar
maxillae, and a labium or lower lip. This basic
segments bearing walking legs. The appendages
design has been highly modified in different groups
are all uniramous (single branched), in contrast to
for acquiring liquid food by piercing, lapping or
the biramous (forked) appendages of crustaceans.
sucking.
An important difference between centipedes and
 48

The thorax is specialized for locomotion, while The sexes are separate, and insects are typically
the abdomen is specialized for digestion, excretion oviparous (egg-laying). Nearly all insects undergo
and reproduction. Each thoracic segment has a pair metamorphosis, with more or less drastic
of jointed legs, and the last two thoracic segments morphological change taking place during
bear a pair of wings. These may be modified in development. The hard exoskeleton restricts
various ways, flies having only one pair of wings and growth, and the insect can increase in size only by
some insects losing both pairs. Wings are not jointed moulting. The new cuticle need not be exactly the
appendages but develop as outgrowths of the body same form as the old, which makes metamorphosis
wall. Flight muscles occupy most of the thorax. possible.
Flight is important in dispersal and in finding food
Insects can be divided into two broad groups
and mates and is a major factor in the success of
according to the extent of their metamorphosis. In
insects.
insects with incomplete metamorphosis the larva,
Respiration in insects: called a nymph, does not differ greatly from the
adult, although it lacks wings. A familiar example is
Instead of oxygen being transported in the blood,
the locust. Complete metamorphosis occurs in
as in most animals possessing blood circulatory
about 80% of insect species, including butterflies,
systems, in insects it reaches the cells of the body by
flies and beetles. Here the larvae look utterly
diffusion through a system of ramifying tubes called
different from the adult and usually have different
tracheae. These open to the exterior by paired
feeding habits. After several larval moults there is a
spiracles. This kind of respiratory system limits the
pupal stage, when the insect does not feed and the
size of insects. They possess a dorsal tubular heart
larval tissues are broken down and adult tissues,
and an open circulatory system, with the blood or
including wings, are formed.
haemolymph directly bathing the tissues. As in all
arthropods, the nervous system consists of a brain Highly developed social behaviour is found in
and double ventral nerve cord with fused ganglia, termites (Isoptera) and in bees, wasps and ants
but the number of ganglia is reduced in the adult. (Hymenoptera). Social insects have a caste (class)
system with division of labour, i.e. dedicated
functions (e.g. queen, drone and worker honeybees),
and colony members feed offspring that are not their
own. Social behaviour depends on complex
communication systems. Some benthic crustaceans,
such as social shrimp, also have a similar complex
social structure.

Subphylum Crustacea
Fig. 9 The respiratory system of insects (Hickman The crustaceans (from the Latin “crustaceus” =
et al. 2012). hard shelled) include such familiar groups as the
 49

lobsters, shrimps, crabs and barnacles, and are the Class Branchiopoda
dominant arthropod group in the sea (where both Branchiopods ("branchi" = gill + "podar" = foot)
chelicerates and hexapods are poorly represented). typically have a series of flattened, leaflike trunk
Crustaceans are also abundant in freshwater, but appendages, which are used for both respiration,
only a few specialized groups, such as land crabs and filter-feeding and locomotion. The trunk segments
woodlice, occur on land. are not fused to the head, but in other respects there
Crustaceans differ from other living arthropods in is considerable variation in body form. The most
two ways: (i) they have two pairs of antennae, familiar representatives are:
whereas all other arthropods have either one pair or  The order Anostraca, including fairy or brine
none; and (ii) their appendages are biramous or shrimps (genus Artemia). These are slow swimming
Yshaped. These appendages consist of a basal and defenceless crustaceans, which lack any
segment (called the propodite), which branches carapace, and have many undifferentiated trunk
distally into two rami, (an inner endopodite and appendages. They swim upside down in temporary
outer exopodite). ponds and salt lakes, forming the diet of many large
Apart from this, one of the main features of the predators, and are well known for their abilities to
Crustacea is their incredible variation in structure. produce egg capsules which can remain dormant (in
There is no such thing as a "typical" crustacean. Even diapause) for up to 10 years. These hatch as soon as
the basic body regions are incredibly variable, with they are wetted and are marketed as live food for
some groups possessing a separate head, thorax and aquarium fish. The red tinge of saltpans at Coega are
abdomen, others a head plus undifferentiated trunk, caused by high densities of Artemia.
or fused cephalothorax plus abdomen, and so on.
 The order Cladocera, including water fleas
The limbs are equally variable and may be adapted
like Daphnia, which are extremely common pond
for walking, swimming, filter feeding, burrowing or
animals. They have short almost circular bodies,
a host of other specialized functions. Indeed, at first
covered by a large carapace and swim using their
sight it is difficult to appreciate that organisms as
enlarged second antennae. Females can reproduce
diverse as, say, an acorn barnacle, a parasitic
parthenogenically (without fertilization) in spring
copepod and a crab, all belong to this single group of
and summer, carrying the eggs in a special brood
animals.
chamber within the carapace. The over-wintering
There are approximately 40 000 species of eggs are generated sexually. Daphnia are also part of
Crustacea. These are divided into six classes, three of the freshwater zooplankton and are fed on by a host
which are (the Cephalocarida, Remipedia and of predators such as small fish, freshwater insects or
Ostracoda) are either rare and/or microscopic and tadpoles.
are not treated further. The other, more familiar,
classes are:
 50

Class Hexanauplia Fig. 10 Life cycle and internal structure of the

barnacle Tetraclita sp. (Branch and Branch 1984).
Subclass Copepoda
In the more familiar, acorn barnacles, the cypris
Copepods ("kope" = oar + "podos" = foot) are
larva glues itself to the rock by its first antennae and
generally minute (< 2 mm) but of the most abundant
secretes a circle of calcareous plates around the
multicellular organisms occurring in most marine
larval carapace. In the process of metamorphosis,
and freshwater systems. They have a cylindrical
the abdomen also becomes reduced, the gut becomes
body usually consisting of a head, a six-segmented
V-shaped and the biramous appendages become
thorax and a five-segmented appendageless
long and feather-like in order to filter food particles
abdomen. Most species are planktonic, swimming
from the water.
with their enlarged first antennae, but many others
are benthic or even parasitic, attaching to hosts As a further adaptation to sessile life barnacles

ranging from sponges to whales. are the only crustaceans, which are hermaphrodite,
although they still prefer to fertilize adjacent
Class Maxillopoda individuals, using the extraordinarily long penis to

Subclass Thecostraca Infraclass Cirripedia accomplish this feat!

(“curl”-footed)
Class Malacostraca
The cirripedes, or barnacles, which are all The Malacostraca ("malakos" = soft + "ostreion" =
marine, are the most highly modified of all Crustacea. shell) is the most diverse class of crustaceans and
The adults, which are either sessile, epibionts, or includes many of the more familiar forms. The best
parasitic, are effectively headless, lack an abdomen known of the orders are included under two
and show little or no evidence of segmentation. superorders:
Indeed, the parasitic forms, Rhizochephala, resemble
fungi, consisting simply of a network of fine tubes Superorder Peracarida

spreading through the tissues of the host (almost  Order Isopoda, are primarily marine but
always a crab) to get nutrients from the host. An include the woodlice and pill bugs, which have
external sac contains the gonads. The true adapted to a terrestrial life style and are important
crustacean origin of these species is really only decomposers of leaf litter. They are dorsoventrally
evident in the larvae, which goes through two typical flattened and have seven similar pairs of walking
planktonic naupliar and cypris stages. legs ("iso" = same, "podos" = foot).

 Order Amphipoda are another largely

marine group which include freshwater and
highshore (beach hoppers) species. They can be
distinguished from the isopods by being laterally
flattened and having the first two walking legs
modified into claws ("amphi" = different, "podos" =
foot).
 51

Superorder Eucarida eyes is present, and these are typically mounted on

movable stalks. There are two or three series of gills,
 Order Euphausiacea, or krill, is a group we
attached to the bases of the thoracic appendages and
will rarely encounter, but it is important in marine
enclosed in a gill chamber, formed by down-growths
food chains. Krill links phytoplankton with
of the carapace. The water in the gill chamber cannot
zooplankton and higher animals such as baleen
be allowed to remain stagnant, as the oxygen it
whales. Krill is also used in aquaculture as animal
contains would soon be used up, thus ventilation of
feed.
the chamber is necessary. This is achieved by means
 Order Decapoda ("deca" = ten, "podos" = of a flap of the second maxilla, called the baler,
foot) are the most diverse of all crustacean orders which fits across the front of the chamber and by
and include the familiar crabs, shrimps and lobsters. moving rhythmically pumps water through it and
Because lobsters are so large and abundant, they are over the gills.
often used to illustrate general crustacean structure
The excretory system consists of a single pair of
and function. However, we don’t have lobsters with
green glands, with ducts opening at the bases of the
large pincers in South Africa, but instead have
second antennae. In females, developing eggs are
species of rock lobster with small pincers (which is
attached to the pleopods, and hatch into planktonic
what you would eat in a South African restaurant).
larvae. In crabs, the abdomen has lost its locomotory
The body of a lobster (or rock lobster) is made up function and has become reduced and folded
of two regions, a cephalothorax, derived from the beneath the thorax. Its only remaining function is in
fusion of the head and thorax, and a muscular "tail" reproduction.
or abdomen. The head consists of six fused segments,
the thorax of eight segments and the abdomen of six
segments. Each segment except the first bears a pair
of appendages and the last segment of the abdomen
has a terminal telson. The head bears the usual
crustacean appendages (two pairs of antennae,
mandibles, two pairs of maxillae). However, their
first three limbs on the thorax are no longer used for
locomotion, but are modified as additional Fig. 11 Cross section of the lobster (Branch and
mouthparts, the maxillipeds. This leaves five pairs Branch 1984).
of thoracic appendages to act as walking legs, the
pereiopods. The abdominal segments each carry a Phylum Onycophora (Velvet worms)
pair of pleopods, used in swimming, except for the These unique animals resemble a cross between
last segment, which has a pair of uropods. an annelid and an arthropod. They do in fact have a

Covering the head and thorax is a rigid plate, the mixture of features from each of these phyla. At first

carapace, so that the thoracic segments can only be glance they look like a “worm” and were in fact first

seen on the ventral surface. A pair of compound classified as a slug. The body is soft and elongate,
 52

with segmentally arranged excretory organs The Onychophora is a small group with only
(nephridia). The body wall is muscular, and the about 70 living species, found in tropical parts of
reproductive tracts are ciliated (cilia are rare or the world and also in some southern zones, such as
absent in arthropods). In spite of these typical South Africa, which has a number of species. Four
annelidan features they have a number of other species occur in the forests of Table Mountain.
characteristics that are associated with the Velvet worms are of special evolutionary
arthropods. Their arthropod features are as follow: importance because they are often considered to be
The first pair of legs has become hardened with an evolutionary link between the annelids and
chitin and forms a pair of jaws. Although apparently arthropods. They appear not to have changed much
soft skinned, velvet worms do have a chitinous since their appearance in the fossil record in the
cuticle, which although protective is highly Cambrian period, 500 million years ago. However,
permeable to water loss. The animals are only able there is debate as to whether they could have given
to survive in very moist forest litter habitats. They rise to the arthropods (especially the
have a large brain, open blood system, and myriapod/insect line), or if they merely
tracheal respiratory system. The large antennae represented a type of animal that was a probable
and eyes are similar to those of both worms and ancestor for the arthropods.
arthropods. Eyes are small and lie at the base of the
A proposed clade, Panarthropoda, is suggested
antennae, and the animals avoid bright light and are
to include Arthropoda, Onycophora and Tardigrada
largely nocturnal. They can have up to 43 pairs of
(or water bears) to highlight their evolutionary and
legs, each of which bears a pair of typical
ecdysozoan similarities. Tardigrades will not be
arthropodan claws at the end. The body is evenly
discussed further in this course.
covered in small warts, which are in turn covered in
small scales. The scales give the colour to the
animals, which can be green, blue or orange (the
local species are brown). All have a velvety The In-Between Phylum
appearance as a result of the scales and the warts.
The phylum Chaetognatha (“Bristle-jaws” or
They are carnivorous and are able to subdue
arrow worms) is an interesting group because they
large prey by squirting them with a very sticky gum,
are animals that were previously included under the
which is ejected from the first pair of legs, the oral
deuterostomes because of their embryonic
papillae. This sticky gum can be squirted for 50 cm
development. However, their evolutionary place is
and hardens when it makes contact with the prey.
still debated, and molecular ecologists consider it
The prey is then cut open with the mandibles. They
part of the protostome clade but not belonging to
have developed a wide array of reproductive
either Lophothrocozoa or Ecdysozoa. It is likely that
methods. Some lay eggs which give rise to live
this may be a common ancestor or a unique body
young (oviparous). Most, however, produce live
plan altogether. However, they are ecologically
young (viviparous).
important, as they are torpedo-shaped, marine
 53

predators living in the plankton. They have a thin

cuticle, complete digestive system, coelom and a
hooded circlet of spines around the mouth. They
possess a nervous system but no excretory system or
respiratory system. All are hermaphrodites.
 54

DEUTEROSTOMIA
Deuterostomes is an entirely different clade of
CLADE AMBULACRARIA
metazoans we have dealt with thus far. Even though
they are all triploblastic coelomate animals, their Phylum Echinodermata
development is different. They undergo radial
(Starfish and their kin)
cleavage (not spiral cleavage) and in the blastula
The echinoderms include starfish, sea urchins,
stage the first opening, the blastopore, develops
brittle stars, feather stars and the sea cucumbers.
into the anus and the second opening becoming the
Most of them share three characteristics: radial
mouth. The coelom also develops through a different
symmetry in the adult stage, spines in their skin,
process; in deuterostomes the coelom forms through
and tube feet.
pouching (enterocoely), as opposed to splitting
(schizocoely) in protostomes with a coelom. Unique features characterise the phylum, and
these are not found in other animals. Typically, the
The deuterostomes are much less diverse than
adults are (penta)radially symmetrical, with five
the protostomes. The first phylum, Echinodermata,
arms (or multiples thereof) stretching out from a
have their symmetry, cephalization and skeletal
central body. They are thus described as being
structure ‘reset’ again to more primitive conditions
“pentaradial”. There is no head or front end, and it
somewhat resembling that of cnidarians. The
is even difficult to decide which surface should be
phylum Echinodermata (starfish and their kin) and
called dorsal and which ventral, because some lie
phylum Hemichordata (Enteroneusta or acorn
with their mouths downwards and others
worms, and Pterobranchia) are grouped under the
uppermost. Curiously, echinoderms begin life as
clade Ambulacraria. The phylum Chordata is the
bilaterally symmetrical, with a pluteus larvae. It has
largest and “newest” group in the deuterostome
a tiny body with long thin lobes covered with cilia to
evolution. This phylum includes the vertebrates.
increase their flotation. As it grows, a special group
of cells is set-aside on the left side of the gut. This
echinus rudiment expands to a miniature version
Fundamental differences between of the adult (which has a radial symmetry), while the
Protostomia and Deuterostomia. rest of the larval tissues are absorbed during
metamorphosis.
Table 1 Comparison between the Protostomia and
Deuterostomia. Bilateral symmetry seems to have many
advantages: locomotion occurs in a fixed direction,
the body can become specialised for ‘forward’
movement, and the sensory organs concentrated on
the head, where they are most needed ‘up front’.

So why should the echinoderms go through this

strange process that converts them back into a
radial symmetry?
 55

Probably because their ancestors were all the oesophagus with diffused nerves (nerve
attached to the substratum, never moved around network) to the rest of the body. Another unique
and were filter feeders. For them, radial symmetry feature is the presence of dermal branchiae or
would have made perfect sense, because they could papulae; these serve the function of skin gills.
use their arms to feed and detect food and threats in
The echinoderms have five different body forms
all directions.
that can be divided into five different classes based
Translated literally, the name Echinodermata on their form and function.
means “spiny-skinned” and refers to the fact that all
members have spines, tiny platelets or spicules Class Crinoidea (feather stars or sea
(called ossicles) in their skins. Some of the classes lilies)
have only minute ossicles; in others they are large
This class contains the most primitive forms of
and fused to form a continuous shell (called a test), echinoderms. It was once a dominant group. The
lying just under the skin. Each of the ossicles is fossil record is rich with crinoids; but nearly all
formed from a single crystal of calcium carbonate
became extinct. All are attached by a stalk or by
and continues to grow throughout the life of the
jointed claw-like cirri. They have a small sac-like
animal, so that even those species that have an
body with the mouth facing upwards, and a short,
encasing shell are not prevented from growing.
coiled gut that ends in an anus placed near to the
Ossicles are formed by the mesoderm, so that they
mouth. All other echinoderms are free-living and
are more equivalent to the bones of vertebrates than
mobile, and most lie with their centrally placed
to the ectodermally derived shells or exoskeletons of
mouths downwards, so they are rather like
other invertebrates that have an external hard upsidedown feather stars. The most obvious feature
encasement. of feather stars is a series of between 5 and 200 long,
Echinoderms are coelomate, and part of the exquisitely beautiful, feathery arms that wave in the
coelom becomes modified into a water vascular water and collect tiny food particles by filterfeeding.
system that forms tubes spreading throughout the
body. These are filled with seawater and serve to Class Asteroidea (the starfish or sea
transport oxygen and dissolved food around the stars)
body. Some of the tubes extend through the body They usually have flattened bodies with five (or
wall and provide a means of locomotion. These tube more) tapering arms that are not obviously
feet can be blown up like tiny thin balloons (the distinguishable from the central portion of the body.
ampulla) by pumping water into them, and then Some, such as the ‘cushion stars’ have such short
controlled by muscles to propel the animal along. arms that they are pentagonal in outline. Starfish can
Echinoderms have no excretory system (and regenerate their arms if one is ripped off: in fact, a
perhaps this is why they are confined to the sea, single arm can regenerate a completely new body;
where osmoregulation is not a problem). The this is a form of asexual reproduction and a new
nervous system is a very simple nerve ring around individual can be generated. However, starfish are
 56

dioeciously (have separate sexes) and reproduce via have no suckers and no longer function in
external fertilization, developing into free- locomotion; they gather food instead. Ambulacral
swimming bipinnarial larvae. The numerous grooves are closed. The mouth lies centrally beneath
ossicles in the skin, makes the starfish spiny or scaly the disc and the madreporite is on the oral surface;
to the touch. Some of these ossicles on the upper there is no anus. Five pairs of bursae (sacks) open on
surface are specialised trident-like jaws the oral surface through bursal slits serving as water
(pedicellariae) that project from the skin and snap circulation openings and discharge of gametes
at creatures that attempt to settle on it. On the lower during reproduction. Gonads are on the coelomic
surface of the body, rows of tube feet line open wall of each bursa. Most brittle stars release sperm
ambulacral grooves that extend from the central and eggs into the water and have with an
mouth (ventral or oral surface) to the tips of the Ophiopluteus larvae. However, a few types brood
arms. These usually end in suckers, for locomotion their young inside their bodies and give birth to
and feeding. A madreporite serves as the opening of miniature adults.
the water vascular system.
Class Echinoidea
Most starfish are carnivorous, using their tube
feet to force open their prey. Others gather decaying (sea urchins, sand dollars and heart
detritus or microscopic algae. The stomach can be urchins)
turned inside out, extruded out of the body and Urchins are usually spherical or pumpkin shaped.
enzymes released externally to digest the food. In A few are very flattened and have become bilateral
some parts of the world, starfish exert an important once again as an adaptation to burrowing in sand
control on their prey species. For example, one like the popular pansy shell or sand dollar. In all
species prevents mussels from taking over the shore species, the ossicles are fused into a hard CaCo3 test
in the western USA. Another, the crown-of-thorns, that covers the whole body and lies just under the
has become famous because it consumes vast skin. When urchins die, their tests wash ashore and
quantities of coral. This threat combined with are popular with beachcombers. From the surface of
climate change might exterminate coral reefs. living urchins, long, mobile spines project and are
their primary defence. Some species have hollow,
Class Ophiuroidea poison-filled spines that break off and inject an
agonising venom. All have pedicellaria (with venom
(brittle stars and basket stars)
glands in some species) that snap at offending
Brittle stars have a flat central disc-like body,
passers-by. The common Cape urchin, Parechinus
with five narrow arms clearly differentiated from
angulosus uses these to paralyse the tube feet of
the disc. The arms are encased in ossicles, but these
starfish attempting to feed on it. Five rows of tube
are arranged so that they articulate and are
feet run from the top down to the ventral mouth, and
therefore very flexible and move in a snake-like
work in conjunction with the moveable spines to
manner to achieve locomotion. The arms break
propel the animal.
easily - hence “brittle star”. Tube feet are present but
 57

Urchins are nearly all grazers. They have five squeezing out and cutting off this part of their guts /
large jaws inside their mouths, supported by discharging Cuvierian tubules (like a lizard breaking
powerful muscles and forming a structure known as off its tail). Others eject long sticky tubules that
Aristotle’s lantern. With this they scrape rock tangle the predator. In the Far East, humans are the
surfaces, removing algae. They are so effective that main predator of sea cucumbers and some species
in many parts of the world they control seaweeds. are sought-after as a delicacy (“beche de mer”).
The common Cape Urchin provides protection to There is an extreme danger of over-exploiting them.
juvenile abalone (perlemoen) and is vital to the In the South Western Indian Ocean, some species are
survival of these commercially important shellfish. heavily exploited, and almost driven to extinction.
Sand dollars and heart urchins feed on fine particles Gas exchange also takes place through the skin. The
deposited in the sand that are collected on podia or madreporite lies freely in the coelom. Tentacles
ciliated tracts. Like other echinoderms, Echinoidea are modified tube feet and trap food via suspension
are also dioeciously, with broadcast spawning, with feeding using mucous. Deep ocean species are
external fertilization forming an Echinopluteus deposit feeders on detritus. Sea cucumbers mostly
larvae. have separate sexes, but some species are
hermaphrodites. Only a single gonad occurs, and
Class Holothuridea (sea cucumbers)
free-swimming larvae are called Auricularia larvae.
Sea cucumbers are the oddest of all the
echinoderms. Their ancestors were pentaradial, but
they have become greatly elongated and sausage
shaped, with the mouth at one end (no
cephalization) and the anus at the other, and they lie
on their sides, so that they have effectively become
bilateral once again. A series of branching tentacles
project from the mouth and gathers detritus or
particles from the water. The ossicles are reduced to
tiny spicules, so the skin is often leathery rather than
spiny. Five rows of tube feet run down the length of
the body. The three lower rows are usually well
developed and provide traction, but the upper rows
often become rudimentary (vestigial) and may serve
a sensory role. Inside the rear-end of the body lie
branching tubules (respiratory tree) that are
connected to the rectum and serve the purpose of
respiration. Weird to think of respiratory water
being pumped in and out of the anus, but that’s what
happens! Some sea cucumbers distract predators by
 58

ANIMALS WITH A NOTOCHORD

BEGINNING OF CHORDATES

Phylum Hemichordata Phylum Chordata

Phylum Chordata refers to animals with a
(Not full Chordates)
notochord at some stage in their life cycle. The
The hemichordates are a small group of
phylum is divided into three subphyla: subphylum
bottomdwelling marine animals formerly
Urochordata (the ascidians, or sea squirts, and their
considered part of the chordates based on the
allies); subphylum Cephalochordata (the
presence of gill slits and a rudimentary notochord,
lancelets); and subphylum Vertebrata (also
but it was subsequently discovered that the
referred to as the Craniata). The urochordates and
“notochord” was in fact an evagination of the mouth
cephalochordates together are known as the
cavity and was not homologous to the chordate
Protochordata.
notochord. Two groups, free-living Class
Enteropneusta (Acorn worms) and colonial Class The five characteristics unique to the chordates are:

Pterobranchia (1-7mm, 30 species) occur. Only one 1) Possession of a notochord (from the Greek
group is common, the acorn worms, which range in “noton” = the back, and Latin “chorda” = a cord): a
size from 20mm to 2.5m and include 75 species. The supportive rod that extends along the length of the
genus animal dorsal to the body cavity and beyond into the
Balanoglossus is found in Langebaan Lagoon on the tail.
SA west coast. They are worm-like but have bodies
2) Pharyngeal gill slits: a series of openings
divided into three regions: an anterior proboscis, a
that penetrate from the anterior part of the gut to the
“collar” and a long trunk that is perforated by two
outside of the body.
rows of gill clefts. They are deposit and suspension
feeders moving food particles by ciliary action on the 3) Tubular nerve cord running dorsally to the
proboscis into the oesophagus. Acorn worms have notochord (and often expanded anteriorly into a
separate sexes and external fertilization. Larvae ore brain).
referred to as Tornaria larvae in some species. 4) A muscular postanal tail (usually used for
Appropriately, the hemichordates (“half swimming).
chordates”) have a three-part coelom and one of the
5) Endostyle which is the precursor to the
five characteristics that distinguish Chordata:
thyroid gland.
pharyngeal clefts. They lack a tail projecting
beyond the anus, have no true notochord, no hollow It is important to note that these features may
dorsal nerve chord and no endostyle, which is the only be present at certain stages in the life cycle.
precursor to the thyroid gland. For example, the notochord, nerve cord and postanal
tail is present only in the free-swimming larvae of
 59

ascidians and are not visible in the adult. Similarly, diameter. The feeding process is assisted by a
the gill slits become closed in most adult vertebrates mucus sheet, which is formed in a ventral ciliated
(except fish) but remain visible during development. groove called the endostyle and glides across the
inside of the pharynx. Food particles become
Of the three subphyla of chordates, only the
trapped in this mucus as it passes over the gill slits
vertebrates have developed bony or cartilaginous
and pass with the mucus into the gut. After
vertebrae to protect the nerve cord and a skull
digestion, the faeces are voided through the anus,
(cranium) to enclose the brain. They are treated in
which empties close to the atrial siphon.
the next section. The two remaining subphyla are
still considered to be “invertebrate chordates” and All tunicates are hermaphrodite. The fertilized
are discussed below. These animals do not possess eggs, which may be retained within the adult body,
a differentiated head, brain, skull, sense organs, develop into larvae, which superficially resemble
limbs or vertebrae. small tadpoles and are consequently called tadpole
larvae. It is in these larvae that we encounter all the

Subphylum Urochordata (Tunicates) essential chordate features. They have a muscular

tail (i.e. an extension of the body behind the anus)
The larvae of these animals possess all the
and a tubular, dorsal nerve cord runs the length of
chordate features but in adults most of the features
the tail and into the head, while below it runs a well-
are lost. The 1400 species of urochordates are
developed notochord. The head also contains a
divided into the following three classes:
pharynx with gill clefts, closely resembling that of
the adult.
Class Ascidiacea
After a brief free-swimming life, the tadpole larva
The ascidians are the largest and most familiar
attaches itself by its head end to the substratum, the
group and contains the true ascidians or sea
tail is resorbed and the internal organs rotate 180°,
squirts. These are sessile marine animals, which
so that the mouth or inhalant siphon points upwards
typically attach their sac-like bodies to rocks,
into the water column and the gut is bent into a U-
pilings, hulls of ships or other solid objects. As
shape.
adults they show few typical chordate
characteristics. There is no sign of a notochord, The body of most tunicates are surrounded by a
dorsal nerve cord or tail and the only recognisable cellulose casing called the test or tunic, hence the
chordate feature which remains is a huge pharynx common name, derived from the Latin “tunicatus”,
or “branchial basket” with gill slits. which means “to wear a tunic or gown”. The larger
species, such as the well-known redbait,
Water is sucked in through an inhalant siphon,
Pyura stolonifera, which can be as large as a rugby
passes through numerous tiny slits in the pharynx
ball, are all solitary. However, many smaller species
and then exits through an exhalant or atrial siphon.
are either clumped (occur attached to one another
The pharynx acts as both a gill for the extraction of
in groups) or colonial / compound, consisting of
oxygen and a feeding organ that can filter out food
large numbers of minute zooids embedded in a
particles as small as 1/1000 of a millimetre in
 60

communal test. The individuals in such colonies secrete an elaborate but temporary gelatinous
reproduce by budding and are often grouped in “house” around themselves and propel water
circles or rows around shared atrial openings. through this by beating of the tail. A screen of fine
fibres across the entrance to the “house” filters out
Class Thaliacea food particles, which are then passed back to the
Class Thaliacea is a small group of about 70 mouth in a mucus film.
planktonic species, which use their feeding
currents as a means of propulsion as well as Subphylum Cephalochordata
feeding. Two different lifestyles occur.
Cephalochordates, or lancelets (e.g. Amphioxus),
In the order Pyrosomida a large number of are small (up to 5 cm long) tadpole-like animals,
colonial zooids line a cylindrical test, which can be which are widely distributed, in clean marine sands.
several metres long and is open at only one end. They are elongate, laterally compressed and nearly
Each zooid is positioned such that its inhalant transparent, but in spite of their streamlined shape
siphon on the outside of the cylinder and its they are relatively weak swimmers, spending most
exhalant or atrial siphon on the inside. All the of their time in a filter-feeding position with the body
feeding currents thus suck from the outside and mostly buried in the sand and just the head showing.
discharge into the central cavity, jet-propelling Water enters the mouth (inhalant siphon), passes
the colony through the water. into the pharynx and through the gill clefts into the

The orders Salpidae and Doliolida, by contrast, atrium and out through an atriopore (equivalent to

are mainly solitary, barrel-shaped organisms and an exhalant siphon). Food particles are trapped by

have their inhalant and atrial openings at opposite mucus in the pharynx and channelled along ciliated

ends of the body. Water currents produced by grooves into the intestine. Gaseous exchange is

muscular contractions of the body wall propel through the general body surface, although the

them through the water. pharynx and gill clefts may become important in
respiration while the animal is buried.
Class Appendicularia In many ways the cephalochordates provide a link
Class Appendicularia (also called Larvaceans) between the invertebrates and the third chordate
are another small planktonic group of only about subphylum, the Vertebrata. The cephalochordates
70 species. They resemble tiny ascidians (5 mm), have a hollow dorsal nerve cord, and a notochord
which have retained the larval tail into adulthood, extending the length of the body, as well as a
complete with its hollow nerve cord and postanal tail. Indeed, at first glance the animal could
notochord. (The retention of juvenile be mistaken for a small fish. The cephalochordates
characteristics, such as a larval tail, in adult stages, differ from the vertebrates, however, in their lack of
is known as neoteny.) The pharynx of larvaceans, a differentiated head, paired sense organs and a
however, only has a single pair of gill clefts and is brain, as well as the absence of limbs, vertebrae and
useless for filter-feeding. Instead, the animals
 61

a skull. Cephalochordates are also the only group to activity levels in vertebrates and larger body sizes.
retain all chordate features into adulthood. Adaptations were also necessary to support the
changed habitat use and lifestyle (like locating,
Subphylum Vertebrata
capturing and digesting food or prey), and metabolic
(Animals with a Backbone) demands of vertebrate life. These adaptations
include:
The Vertebrata have two major advances on the
Protochordata. Firstly, the possession of a skull (or 1. Musculoskeletal modifications: strength for
cranium) of cartilage or bone, which protects the movement and protection e.g. change in muscle
brain. The second major advance is they have structure (from V to W shape) and cartilage
vertebrae which are segmented, and which changed to bone.
surround and replace the notochord. These
2. Physiological changes to the circulatory,
vertebrae, which may be constructed of either
digestive and respiratory systems. to support
cartilage or bone, provide more rigid, and thus
metabolism and food processing. The branchial
better, support to the dorsal nerve cord than did the
basket used for feeding shifted to be exclusively
notochord found in the Protochordata. However, the
used for respiration, and cilia for feeding became
development of a skull was preceded with one very
redundant as large amounts of large food
important evolutionary event; the modification of
particles were ingested. Liver and pancreas
first pharyngeal arch support structures to form a
became important to assist with digestion as did
jaw. This enabled a much larger, much more effective
haemoglobin in erythrocytes for increased
method of feeding. Animals in the Vertebrata
oxygen demands.
(Craniata) also contain paired eyes, and other
sensory structures on the head. Another adaptation 3. Enlarged head and brains developed because

unique to gnathostomes, is paired pelvic and of better-developed sensory systems (e.g.

pectoral girdles (seen from the earliest fossil eyes with lenses, olfactory senses, inner ear for

evidence in Myllokunmingia). balance and pressure, lateral line for vibrations)

all used for prey location and capture.
Adaptations:
4. Hox genes (which codes for the body plan), most
There are nine classes within the subphylum
likely coded for two new innovations, the neural
Vertebrata, five of which are fishes. These five
crest (ectodermal cells on the neural tube coding
classes of fishes contain more than a half of all known
for structures like the cranium and pharyngeal
vertebrate species. Craniates includes fishes,
skeleton, tooth dentine) and ectodermal
amphibians, reptiles, birds, and mammals. Hagfishes
placodes (embryonic tissues giving rise to most
are the only modern craniates that are not strictly
of sensory structures) of the head respectively.
vertebrates as they possess no vertebrae in any
These modifications were most likely possible
form.
because most living gnathostomes have four sets
Adaptations that have given rise to vertebrate of Hox genes as opposed to one in
evolution have been advantages to support higher cephalochordates.
 62

Other specific evolutionary advances that we vertebral column consists of two major parts,
should recognise across the evolution of tetrapods including a notochord, and a fibrous tube around
include the modifications necessary to from living in the nerve cord. This shows the beginnings of
water to movement onto land. Fish have a single vertebral development.
blood circuit from the heart to gills (respiratory
Agnathans have pseudo-teeth composed of
organs in water) to the body. However, with the
keratin attached to the tongue, orientated in rows.
development of lungs, a double respiratory system
They are mostly parasitic, scavenging or predating
(two circuits) was more efficient. Kidneys adapted to
on small prey. The metabolism is comparatively
conserve water, and internal fertilization was more
slow (to other vertebrates) with no distinct stomach,
appropriate. And the skeletal structure was
but rather an undifferentiated long gut,
realigned to deal with gravity because the body was
homogenous throughout the body. Agnathans are
no longer supported in water.
dioecious, with external fertilization.
FISHES
There are more than 80 species of extant
A recent fossil discovery, date early fish evolution agnathans, into two classes; Myxini or hagfishes
back to 530 MY ago with the finding of and Petromyzontida the lampreys. Hagfishes are
Myllokunmingia. There are many examples of early exclusively marine creatures. They are predators of
fishes but few of them survived. One example invertebrates, but also scavenge dead material
includes the Ostracoderms. They form an extinct from the seabed. They have a unique feeding
assemblage of jawless fishes recognised by their mechanism that uses leverage against prey body by
protective shell-plated skin (“ostracon” + “derma”). tying their own body into a knot and pulling head
Extant jawless fishes are grouped under a single backwards through the knot. These fishes possess
superclass Agnatha but are very different from each 90-200 slime glands and secrete vast amounts of
other. slime as a predator avoidance mechanism to
disorientate and suffocate predators. Lampreys
Superclass Agnatha (Jawless fishes) have both freshwater and marine representatives.

Phylum Chordata They are all ectoparasitic on other fishes using an

anticoagulant to draw fluids from host.
Subphylum Vertebrata

Superclass Agnatha Superclass Gnathostomata

In general, the Agnatha are jawless fishes, which The gnathostomes include all of the jawed fishes
exhibit the early stages of development of vertebrate (bone or cartilage), as well as all the tetrapods
features. A skull is present which, like the rest of the (four-legged creatures). This superclass includes the
skeleton, is made up of cartilage, a precursor to extinct placoderms; cartilaginous fishes
bone. Unlike more advanced fishes, there are no (Chondrichthyes) and bony fishes
paired fins, only median fins that run down the (“Osteichthyes”). The Chondrichthyes are fish
mid-line of the dorsal and ventral body surfaces. The with a cartilage skeleton, gill slits, and teeth that
 63

shed and constitute sharks, rays and chimaeras which completely surround the dorsal nerve cord.
whereas bony fishes (“Osteichthyes”) all have a Apart from this, the Chondrichthyes have other
bony skeleton, gill covers, and a swim bladder. They skeletal elements, including the two limb girdles,
can be further separated into the ray-finned fishes which provide muscular attachment and a system of
(Actinopterygii) and the lobe-finned fishes levers for the attachment and movement of the
(Sarcopterygii) like the lungfishes and coelacanth. paired fins. The paired fins are used for a number of
purposes in the Chondrichthyes; In sharks they are
Class Placodermi  used primarily to control rolling of the body from

The placoderms are the armoured fishes, all of side to side and as hydrofoils to direct the shark
which are extinct. They had primitive jaws, paired upwards or downwards in the water. In skates and

fins, and a notochord surrounded by bony rays, the pectoral fins are greatly enlarged, and are

elements. The head and much of the trunk was used almost as "wings" to create lift and forwards

armoured by bony plates formed by the skin propulsion of the animal through the water.

(dermal bone). Respiration in sharks takes place through the 5 - 7

gill slits, and they need to move for oxygenated water
Class Chondrichthyes to flow through gills. Rays and skates, however, have

Are recognised by their cartilaginous skeleton, 5 to 6 gills on the underside of the body and can
actively pump water through spiracles and out of
and a lack of swim bladder. Most store oils and
ammonia in the liver to assist with buoyancy. Like gills. Larger shark species give birth to live young

other fishes, sharks have a two-chambered heart while smaller species, rays and skates lay egg cases

with blood flowing between the gills and the body. with external embryonic development.

The shark-shape is typically recognised by Cartilaginous fishes display a variety of feeding

heterocercal tail (sharks and chimeras), and the modes although almost all sharks are carnivorous,
sand-papery placoid scales covering shark skin. feeding on live vertebrate and invertebrate prey.
Most sharks copulate, with internal fertilisation, Some exceptions, like basking and whale sharks are
but they display all forms of embryonic development plankton feeders. Very few sharks are scavengers.
i.e. viviparous, some oviparous, and some
Subclass Holocephali (Chimeras &
ovoviviparous.
ratfishes)

Subclass Elasmobranchii (Sharks & rays) This is a small class of cartilaginous fish of which

This group comprise the sharks, skates and rays. the whole upper jaw is fused to the skull. Chimeras

This group contains roughly 850 species, all of which characteristically have four sets of gill slits protected

are marine, though some (like the Zambezi Shark) with a soft gill cover and only one slit open to the

may enter estuaries and rivers on occasion. Like the outside. They are exclusively marine and tend to

agnathans, they do not have true bone, and their occur in deeper water. They propel themselves with

skeletons are composed of cartilage. The vertebrae large fan-shaped pectoral fins. Common in South

are segmental and are well-developed structures, Africa is the elephant fish or St Joseph ‘shark’
 64

Callorhinchus capensis. They are dioecious, and after

copulation distinctive spindle shaped, egg cases are
laid from which the juvenile will hatch.

THE BONY FISHES:

Superclass “Osteichthyes”

Old classifications refer to the class Osteichthyes

as the bony fishes, because they possess true bone,
but this has since been split into two classes; the
rayfinned and the lobed-fin fishes. Characteristic of
bony fishes is the well-developed skull and
vertebrae. Like sharks, they have paired fins, but
also a swim bladder (usually) to maintain neutral
buoyancy; the fish thus neither floats nor sinks but
can remain at a particular depth in the water column.
Respiration is through the gills covered with by
opercula, with the exception of the lung fishes which
are also equipped with a “lung” for gas exchange. The
circulatory system is the same as all other fishes;
closed circulatory with a two-chambered heart.

Reproduction in body fishes vary; they can be

dioecious or hermaphroditic but have external
fertilization. They are egg-laying, either as
oviparous or as ovoviviparous breeders.
 65

The coelomate condition

It is likely that the coelomate condition has arisen

at least twice (maybe three times), because the
arrangement of body cavities in molluscs and
arthropods is distinctly different from that in the
other coelomate phyla. The embryological
development of both these taxa shows them to be
coelomate, and the gut is surrounded by mesoderm
as it is in other coelomate groups, but a further
complication arises. The coelom is reduced to small
spaces around the heart and a few other organs, and
the major body cavity is a separate entity known as
a haemocoel. While animals with a coelom generally
have two separate systems of body fluids (the
coelomic fluid contained in the coelom and the blood
contained in a closed blood system), those with a
haemocoel tend to have only one. The haemocoel
forms part of an open blood system, the heart
pumping the blood through large blood vessels that
open straight into the haemocoel. In both molluscs
and arthropods, the fluid in the haemocoel may act
as a hydrostatic skeleton.

Fig. 12 Schematic drawing of an open and closed

circulatory system. The open circulatory system is
associated with a haemocoel and the closed
circulatory system with a coelom.
 66

zoological world by the discovery of the coelacanth

(Latimeria chalumnae) near East London, and for
Class Actinopterygii
many years this fish was dubbed "Old fourlegs", as it
The ray-finned fishes are the most common was held up as the ancestor of land tetrapods.
fishes and also the main group consumed by However, it is now well established that the
humans. The paired fins have relatively fragile coelacanths are not the direct ancestors of land
structures of bone, and not have a fleshy base like the vertebrates. Instead, the ancestors are more likely to
lobed-fin fishes. The diversity within the ray-finned have come from a group of extinct fishes called the
fishes is staggering, with more than 24 000 species Rhipidistia, which more closely resembled present
known to science. day lungfishes.
Osmoregulation in fish is achieved through an Lung fishes have gills and a lung-like sac for
interaction of the gills, intestine, and kidneys, which breathing air. They are freshwater species which are
depends on if the fish is in fresh water or sea water. used to ephemeral water sources; Australian species
Both environments are metabolically stressful as it uses gills and cannot survive long out of water,
requires maintenance of osmotic balance against a whereas African and South America lungfish can live
gradient. In freshwater, the fish blood is more out of water for long periods. Aestivation (period of
concentrated (hyper-osmotic) to the environment, dormancy) is via a slime cocoon during dry periods.
so water enters into the body through osmosis. Salt
Coelacanths are the only modern lobe-finned
is lost via diffusion, and dilute urine is produced. The
fish. They arose in the Devonian period and were
opposite is true for fish in sea water. Fish blood is
previously thought to have gone extinct 70 million
less concentrated (hypo-osmotic) relative to sea
years ago (mya).
water. Water is thus lost to the environment via
osmosis and salts are gained via diffusion producing
concentrated urine.

Reproduction in bony fish is mostly oviparous THE TETRAPODS: INVASION OF LAND

with a larval phase and a gradual metamorphosis. A
small fraction of bony fish is viviparous, and some Class Amphibia (Frogs, newts and
like the klipfish (Clinidae), are ovoviparous. salamanders)
Terrestrial vertebrates appeared in the late
Class Sarcopterygii Devonian (~400 mya), a time when terrestrial
The lobe-finned fishes are an interesting group ecosystems comprised tropical ‘forests’ of ferns and
and include the coelacanths and the lungfishes. The horsetails swarming with large insects and other
bases of their paired fins are thick and fleshy, and land arthropods. The first vertebrates to invade land
almost limb-like. Indeed, lungfishes use these were primitive amphibians, and these had to face
successfully to crawl over land. It is thought that the numerous challenges that the new terrestrial mode
ancestor of land vertebrates was a kind of lobe- of life imposed: the reason for this is that almost all
finned fish. Great excitement was caused in the biological systems operate very differently in air and
 67

in water. Aquatic animals are supported by water, These early limbs displayed a pattern, which
but sturdy skeletons and limbs are required on land. persists today in all vertebrates – the presence of
Gills, which extracted oxygen from a unidirectional five digits in the feet and hands (the pentadactyl
flow of water, have had to change to large volume, condition). Most amphibians have four front toes
thin-walled sacs - the lungs which moved air in and (“fingers”) only though. The type of limb present in
out of structures. Apart from numerous other Ichthyostega is still found in living examples of
adaptations which were required to avoid the primitive fish, and it is these fish (or their close
possibility of drying out (not only of the adult stage relatives) that were the probable ancestors of early
but also the eggs), there had to be additional changes amphibians. Most of them are air breathing
in the sensory systems which function very freshwater inhabitants. Recent fossil evidence has
differently in air and in water. New sensory channels shown another intermediate animal between lobefin
such as hearing were developed, while vision was fishes and tetrapods called Tiktaalik. Fossilized
also improved. Such changes that were initiated with specimens of this animal is dated to 375 mya.
the Amphibia have been continued up the line of
Many of the changes that occurred in the
vertebrate evolution.
transition from water to land were internal. Some
At some evolutionary stage the first amphibian of the more obvious changes were those that
ancestor must have had a mixture of fish and occurred in the circulatory and respiratory
amphibian features. Among the very large fish-like systems. The Amphibia are unique amongst the
animals that are found in the fossil record just before vertebrates in that they are able to extract a fair
the appearance of early Amphibia, there is one that amount of oxygen from the air through their skin. It
fits all the requirements of an ancestor of the first is generally moist, and well supplied with blood
terrestrial vertebrates, a fish-like amphibian. vessels. Some have a glandular skin, used for
Ichthyostega was an inhabitant of freshwater ponds defence as it secretes poisons. In common with all
and had a complete mixture of fish and amphibian respiratory systems it also serves for excretion of
characteristics, indicating that although it spent carbon dioxide. However, active vertebrates cannot
some time in the water, it must have been capable of rely on this passive method of gaseous exchange to
dragging itself out of the water at the margins of the obtain sufficient oxygen. Thus, all land vertebrates
pond, and breathing air. We know that it was have developed changes in the heart and associated
airbreathing, as it had nostrils that led to lungs. blood vessels to pump blood at a faster rate to
The limbs were modified so that they could support oxygen starved tissues. These changes in the
the body out of water: the forelimbs were much circulatory system also move the deoxygenated
stronger than the hind limbs, and the muscles of the blood to a pair of new organs, the lungs. These sacs
limbs were positioned outside the body. The bones, have a large surface area. In the Amphibia, the
which supported the ray fins of fish had now moved heart is much larger than that of fish, and has been
outside the body, were reduced in number and divided into three chambers, separating
strengthened, forming the bones of the limbs. oxygenated and deoxygenated blood. This is the first
step in the evolution of the very efficient
 68

fourchambered heart and associated double (metamorphosis).

circulatory system characteristic of higher
There are three orders of which only one, the
vertebrates. In Amphibia, oxygenated blood
frogs and toads, occurs in Africa. The two other
reaches the heart via the pulmonary vein. It enters
orders of living Amphibia are the salamanders and
one of the atria, while deoxygenated blood that has
newts, and the caecilians.
been drained from the rest of the body enters
through the other atrium. At this stage, the two Order Anura (Frogs and Toads)
kinds of blood are kept separate. Although they then They all have a short body and enlarged hind
enter the single ventricle there are a number of limbs for jumping. The tadpoles are typically
adaptations that do in fact keep the blood types aquatic, but many species have been able to breed
relatively separate. Thus, the more oxygenated independently of water, with the tadpole stage being
blood is pumped to the head and anterior parts of suppressed in the few large yolky eggs that are laid
the body, while predominantly deoxygenated blood in nests in the soil. These eggs hatch into small
is pumped to the skin, where carbon dioxide is froglets. Adult frogs are all carnivorous, while the
‘breathed off’, and oxygen loaded. If the oxygenated tadpoles are generally algal feeders. The transition
blood went directly from the lungs to the tissues, between tadpole and the adult frog is dramatic, and
there would be a large pressure drop, as the blood termed metamorphosis. Many body parts have to
must cross two capillary beds in the lungs. To give be broken down and replaced by new organs that
the blood a boost in pressure it is directed back to suit the adult frog’s lifestyle. The skin is very
the heart via the pulmonary vein, and then shunted glandular, with mucus glands helping to keep the
to organs and tissues under high pressure. skin moist and alkaline glands producing toxins, that
in some frogs, such as the poison arrow frogs, are
deadly in minute amounts. Frogs also have more
LIVING AMPHIBIA
colour producing cells than any other vertebrate,
Characteristic features of living amphibia are that
and many of the poison arrow frogs are very brightly
they have a hollow, ossified bony skeleton but the
coloured. The ear is well developed, as frogs make
body shape varies greatly among orders. They are all
extensive use of sound to locate mates. However,
ectothermic (so the body temperature is modulated
there is no external ear (pinna), and the eardrum
by external environment). They all have a scaleless,
of the middle ear can generally be seen on the
moist skin, for respiration. The skin can also be outside of the head.
glandular secreting chemicals for defence. They are
There are more than 5000 species of anurans
mostly four-legged; their forelimbs generally have
known from 251 mya in Triassic. Amphibians with
four digits. The heart is modified to a
four legs, with long, muscular hind legs. Tadpole
threechambered heart with a double circulatory
(larvae) have tails which are lost as they
system between the lungs and the head (via the
metamorphose to the adult form. They are all
heart), and the skin and the heart. Amphibians are
ectothermic frequenting aquatic habitats excluding
dioecious, with direct or indirect development
the extremely cold polar regions. Toads are
 69

generally thick-skinned, specialised to terrestrial Order Urodela (Salamanders and Newts)

habitats. They are still tied to water for reproduction.
Salamanders and newts are common in northern
Anurans use cutaneous respiration while submerged
temperate regions in Eurasia and the Americas with
underwater and lungs on land.
553 species. As they have far more vertebrae than
Most frogs hibernate in winter.
frogs and appear longer bodied, with tails. In both
External fertilisation (AMPLEXUS) groups the larvae have external gills and are

Amphibians depend on standing water to breed. aquatic, but in the salamanders the adults lose gills

It seems like a simple strategy, but these features and move into dry land. In the case of the newts the
adults are also totally aquatic. All use chemical cues
make them vulnerable in the Anthropocene to
for mate location as opposed to sound. They vary in
habitat loss (infill of wetlands), disease, and
size, but the giant salamander of Asia reaches almost
pollution (endocrine disrupters) causing deformities
two metres in length!
and threatening species.
Body form most like early tetrapods, side-to-side
Order Gymnophiona (Caecilians)
walking motion, with rudimentary limbs,
These are worm-like burrowers, which
sometimes absent. Some are burrowing but mostly
resemble large earthworms more than amphibians.
aquatic and in newts they may retain gills as adults
They are totally terrestrial, inhabiting the moist leaf
(neoteny or paedomorphosis). They are
litter of tropical forests. They have low diversity,
carnivorous feeding on worms, arthropods,
with ~170 species which includes many limbless,
molluscs.
blind burrowers, and some aquatic species,
Reproduction starts with internal fertilization.
inhabiting tropical forests in South America, Africa,
In terrestrial environments, the female collects a
India, Southeast Asia. Like frogs the tail is absent or
sperm packet deposited by male on leaves. Egg
short if present.
clusters are laid under logs or in damp soil with some
Some have small dermal scales on skin, eyes
species guarding eggs or they have direct
small or many are totally blind as adults, as eyes are
development. In aquatic habitats females lay eggs in
redundant underground. They eat worms and small
clusters which develop into an aquatic larval phase.
invertebrates.
Some newts have double metamorphosis; first from
Reproduction is functional to the environment water to a land juvenile form and then to an aquatic
with internal fertilization using protrusible adult.
copulatory organs in males. Females deposit eggs in
moist ground near water or guard the eggs in body Class Reptilia (turtles, snakes, crocodiles)
folds in some species. Eggs hatch as free-swimming Rise of the Amniotes: Amniotes are animals with
larvae, when in water or with larval development in embryos that develop inside a waterproof egg.
egg. A few are live bearing (viviparous). They have waterproof skins (of keratin and lipids)
and highly efficient kidneys make them well adapted
to dry habitats on land. Reptiles are paraphyletic
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group, so an artificial group referring to amniotes sensory abilities and coordination of

excluding the bird and mammals. locomotion.
Dinosaurs are extinct amniotes with birds as their
In the late Carboniferous, about 300 million
descendants, and mammals started from the
years ago (mya), amphibians still dominated most
mammal-like reptiles.
land-based niches. Reptiles had evolved some 50
Adaptations of Amniotes: mya earlier from amphibian stock but they were
small, lizard-sized tetrapods. However, by the
1. Amniotic egg with three egg
Permian era (about 286 mya), reptiles became the
membranes: the amnion, allantois, and
dominant land vertebrates. Their success was
chorion, and nutrients contained in a yolk sac.
largely due to their complete independence from
2. Thick waterproof skin with hard water. They achieved this specifically through the
(beta)keratin and hydrophobic lipids for following adaptations: they abandoned respiration
protection against desiccation. through the skin, which became dry and covered
3. Ventilation of lungs using ribs, with scales. Water loss through the skin was thus
instead of mouth or pharyngeal pressure like greatly reduced. This only possible as the lungs
amphibians. became more efficient and able to supply all the
needs of gaseous exchange. The eggs with semi-
4. Suction feeding in fish is replaced
permeable shells were laid on land. This cleidoic
with a stronger jaw with teeth, and a tongue,
(closed box) egg contained enough nutrition for the
allowing mastication (they chew their food).
embryo to develop into a terrestrial hatchling.
5. High pressure cardiovascular Larval stages were suppressed. The excretory
system with three heart chambers; an atrium products of the developing reptile (now mainly uric
that receives deoxygenated blood from the body; acid) were stored and simply left behind when the
and an atrium that receives oxygenated blood egg hatched.
from the lungs. Both drain into a ventricle which
The major reptilian lineages are separated on the
is partly separated. In diving animals (like sea
basis of the number of temporal openings in the
turtles) the blood is shunted away from the
skull. The anapsids, which includes the turtles, do
pulmonary system during diving, as animals are
not have any temporal openings. The synapsids,
not breathing anyway. Crocodiles, however, have
including the mammal-like reptiles, have a single
a four-chambered heart, like birds and mammals.
pair of openings. Dinosaurs, crocodiles and snakes
6. Water-conserving nitrogen (as well as birds) have two pairs of temporal
excretion in the form of urea or uric acid as openings and are termed diapsids. A fourth group of
opposed to ammonia in most fish. extinct reptiles, which included the marine

7. Expanded brain (cerebrum and ichthyosaurs were the eurapsids; they had a single

cerebellum) which enlarged due to increased pair of temporal openings, but these differed from
those of the synapsids in being surrounded by a
 71

different set of bones. The temporal openings may K-T asteroid impact hypothesis:
have reduced the weight of the skull, conserved
A thin layer of iridium, rare on earth but common
calcium and accommodated the larger jaw
in space (asteroids), and the coordinated sudden
musculature.
extinction of a large fraction of life on earth provide
About 250 mya, South Africa emerged from the support for the Asteroid Hypothesis. Further, a large
grips of a great ice age. As the ice sheets retreated, asteroid site found in Mexican, Yucatan Peninsula
the low-lying, central Karoo basin was revealed. This suggest that it was likely. It was probably followed
became the scene for 50 million years of by massive volcanic eruptions (Deccan Plateau of
evolutionary history. Several groups of vertebrates India) and climate change. Most dinosaurs became
are represented in the Karoo fossils but the most extinct at this time, with a few exceptions like the
famous are those of reptiles and in particular the large leatherback sea turtle.
mammal-like reptiles (synapsids). At the end of the
Today, only four orders of living reptiles remain.
Permian (about 245 mya), a mass extinction event
They are the Testudines (turtles and tortoises), the
occurred and 37 families of tetrapods were reduced
Squamata (lizards and snakes), Sphenodonta
to ten. This extinction affected both carnivores and
(Tuataras) and the Crocodilia (crocodiles). Birds
herbivores on land and 50% of marine invertebrates.
probably arose from small archosaurians, and the
Some reptile groups went extinct, while others were
crocodiles are their closets living relatives. There
greatly reduced in numbers and then became
are altogether about 5 500 living reptile species.
extinct. The Cynodontia arose in the Triassic and
General features of reptiles include scale-
became the dominant mammal-like reptiles.
covered body with beta-keratin and hydrophobic
The dinosaurs evolved in the late Triassic (about
lipids. A cloaca (combined excretory and
225 mya) and for the next 160 million years
reproductive opening) for excretion of uric acid.
dominated almost every land niche. Mammals were
The bladder reabsorbs water from urine to produce
around but were small, probably nocturnal
uric acid. They have four, approximately equal
insectivores. Dinosaurs are divided into two groups
limbs except in snakes, with only remnants of the
based on the arrangement of the pelvic girdle; the
pelvic and pectoral girdle. The skull includes a jaw
reptile-hipped dinosaurs (saurischians) and the
and single occipital condyle (bony protuberance)
bird-hipped dinosaurs (ornithiscians), although the
probably for muscle attachment. Circulation is
birds did not originate from this group, but from the
attained through a three-chambered heart except
saurichians. The saurischians included the
in crocodiles with four hearts. Reptiles have good
meateating dinosaurs and the long-necked,
vision, including colour vision. Some, snakes and
herbivorous dinosaurs. The ornithiscians included a
lizards have a Jacobson’s organ for olfactory sense
variety of plant-eating forms. At the end of the
or enhance taste.
Cretaceous (65 mya), another major extinction
Reptiles are dioecious, with internal fertilization
event occurred, culminating in the extinction of the
(using a penis). All forms of development, i.e.
dinosaurs.
oviparity, viviparity and ovoviviparity take place,
 72

with parthenogenesis in some lizards. Body roof of mouth. Pythons, boas and pit vipers have heat
temperature is dependent on the surroundings and sensing pit organs instead.
are thus ectotherms.
Order Sphenodonta (Tuataras)
Order Testudines (turtles and tortoises)
Tuataras live on rocky islands off the coast of New
Tortoises are terrestrial and turtles aquatic. They Zealand are the sole survivors of the ancient
don’t have teeth but a strong bony jaw. Tortoises are sphenodontid linage. They burrow in (bird) petrel
herbivores but marine turtles feed on a variety of burrows, often sharing the nests with birds. They are
prey depending on the species. Testudines breathe carnivorous feeding on bird eggs, chicks,
through abdominal and pectoral muscle movement amphibians and other small reptiles. They are slow
as they have no diaphragm. They have poor hearing growing and reach sexual maturity between 10-20
and are mute (producing no sounds) other than years. Eggs are slow to incubate which be up to seven
grunts. Like many fish they have environmental sex months, and individuals may live longer than 100
determination so no sex chromosomes, but the years.
incubation temperature determines the sex of eggs
The two remaining species of tuataras have a
(oviparous). Low temperatures produce males and
median parietal “third eye” under the skin of the
higher temperatures females.
forehead, with a photoreceptive function
associated with the pineal gland, regulating
Order Squamata (lizards and snakes)
circadian rhythmicity. Diapsid skull similar to
Most of the non-avian reptiles are included under
specimens from 200 mya.
the Squamata which include the lizards and snakes.
Although well adapted to terrestrial life, they also Order Crocodilia (Crocodiles)
have freshwater and marine representatives (e.g. sea
Only survivors with birds of the archosaurian
snakes). They can be fossorial (live underground) or
lineage. They are semi-aquatic predators, adapted
arboreal (live above ground). The skull is kinetic,
to water living, using a flap of skin (complete
with movable joints, located at quadrate and secondary palate) that prevents water from
pterygoid bones; the snout and upper jaw moves on entering their throats while attacking prey
rest of skull which allow them to capture and eat underwater. They can still breath with a mouth full
larger prey.
of food, like mammals.
Two suborders exist, the Sauria (Lizards) and Crocodiles guard their nests and display
Serpentes (snakes). Lizards have movable eyelids
parental care. They also have temperature
and external ear openings, whereas snakes are
dependent sex determination with temperature
legless, but some have bony remnants of hindlimbs.
ranges opposite to sea turtles, i.e. high temperatures
They gave no external ear openings, no eyelids, with
produce males and low temperatures females.
hearing sensitive to low frequency vibrations. They
Crocodilians are close relatives of birds, and the only
have forked tongues which collects scent molecules,
reptiles with a four-chambered heart.
which are then drawn past the Jacobson’s organs in
 73

Class Aves (Birds) adaptation for an ectothermic animal, as they would

There is little doubt in the minds of most slow down that rate of heat gain. Feathers consist of

palaeontologists that the origins of birds lie among a hollow shaft (calamus) topped with downy

the dinosaurs, probably among the theropods. Until feathers and then followed with a central rachis and

recently, Bavarian specimens of Archaeopteryx vane, the latter consisting of barbs with barbules.

lithographica, some 147 million years old, formed Flight is energetically expensive, with a high
the focus of the reptile-bird transition debate. Recent oxygen demand. Birds have a unique respiratory
discoveries in China (Caudipteryx and system incorporating sponge-like lungs and hollow
Protarchaeopteryx) have provided evidence for the air sacs that are connected to pneumatised bones.
transition from rudimentary filamentous feathers to The volume of the respiratory system is much larger
the streamlined feathering of modern birds. than that of an equivalently-sized mammal. The

Modern birds consist of Paleognathae which are spongy lungs and air sacs are arranged to provide a

ancient birds (such as kiwis, emus, cassowaries and unidirectional airflow through the lungs, which in

ostriches), whereas the rest of the more recent birds turn, allows for a counter-current system

comprise the Neognathae. Birds range in size from analogous to the flow of water over the gills of a fish.

hummingbirds at 1.8 g to the common ostrich that Oxygen extraction is highly efficient and allows birds

weighs 100 kg. The extinct elephant bird is said to to function at higher altitudes than mammals.

have weighted 450 kg. Birds are either passerine Designed to meet high metabolic demands of flight.

(perching) or non-passerine (non-perching). Lungs have parabronchi (not alveoli like mammals)
receive continues air flow. Nine interconnected
The aerodynamic demands of flight have
pairs of air sacs are used in two respiratory cycles
imposed design constraints on birds, including total
for a single breath to pass through respiratory
commitment of the forelimb to flight. Some
system.
reptilian features have been retained, including the
cleidoic egg, but others have been greatly modified. The sensory system of birds is highly developed.

There has been substantial reduction and fusion of This is well reflected in the structure of the eye;

bones to provide skeletal lightness and rigidity; almost the entire image received is in focus and birds

enamelled teeth have been lost and the long bony can change focus (accommodate) very much more

reptilian tail has been reduced to a stumpy rapidly than mammals. The importance of vision to

pygostyle that provides a site of attachment for the birds is highlighted by the ostrich Struthio camelus,

tail feathers. Some modern birds, such as penguins, which has the largest eye of any terrestrial

are flightless, but all evolved from flying ancestors. vertebrate.

Feathers provide excellent insulation, allowing The ability to fly has promoted radiation and

them to maintain a fairly constant, high body speciation. There are approximately 9700 species

temperature (endothermy). It is likely that the of bird worldwide, more than twice the total

immediate ancestors of birds were at least partially mammal diversity. Interestingly, among the

endothermic: insulating feathers would be a poor mammals, some 25% of species are bats, the only
 74

group of mammals that has mastered powered flight. the species, some chicks are precocial (fast
Birds are found in almost every habitat on Earth, developers) whereas others are altricial.
from the open oceans to the poles and the driest
Known bird egg anatomy: The yolk sac provides
deserts. Bird feet and beak design is highly adapted
nourishment for the developing embryo, and the
to meet energy demands of the species. This includes
amnion provides a fluid support and moisture for
swimming, walking, perching, seizing prey or
the embryo. The chorion serves a gas exchange
climbing. Feeding included filtering, probing,
function while the allantois stored waste from
catching insects, cracking seeds, tearing meat
metabolic processes. These parts are all surrounded
and drilling holes. Diets are avivorous (bird eats
by protective albumin and a hardened shell.
other birds), carnivorous (meat eating),
Flight has also opened the doorway to migration.
insectivorous (insect eating), ophiophagous
Although many animals migrate, the migrations of
(specializing in eating snakes), frugivorous birds
birds are the most spectacular of all. Many bird
(eat fruit), granivorous (eat grain or seeds),
species breed at Arctic latitudes during the brief
molluscivorous bird (feed on molluscs), mucivorous
northern summer and then fly south to the Southern
birds (feed on the mucus of plants), nectivorous
Hemisphere to exploit the seasonal bounty of food in
(feeds on flower nectar). Piscivorous birds feed on a
the southern summer. The Arctic Tern Sterna
diet of fish, and omnivorous birds feed on a mixed
paradisaea has the longest migration of all. Some
diet of whatever they can find.
individuals travel from breeding grounds in the
Reproduction in birds is under severe sexual
Canadian Arctic, across the Atlantic, down the coast
selection resulting in sexual dimorphism (male and
of West Africa and around the Cape of Good Hope
female look different like peacocks); Males engage in
before turning east towards their final destination in
courtship displays changing feathers to be overly
Subantarctic seas to the south of Australia. This
long or colourful. (The testes also enlarge in
journey ensures that they see more daylight in a year
breeding season). Most birds (> 90 %) are
than any other animal.
monogamous for life, although extrapair copulation
General features of birds:
is quite common in these relationships. Some are
polygamous. Polygyny also exist with one male Birds are bipedal, egg laying, most nesting with
with a harem of females, frequently in a lek. brooding of eggs. They are feathered for insulation
Polyandry (where one female has many males and and flight. The wings are modified forelimbs
the male incubates the eggs) is a rare condition. adapted for flying or swimming, with a lightweight,
Sexual reproduction, internal fertilization. Most porous skeleton, with strong muscles. These
male birds lack a penis and use the cloaca for sperm animals produce body heat (making the,
transfer (However, ducks have a long corkscrew endotherms). They possess specialized digestive
penis). Most birds lay eggs in nests, some lay eggs in systems with a crop, no teeth, but with beaks very
nest of other species (Cuckoo). Chicks are cared for well designed and adapted to their respective diets.
by one or both parents, mostly both. Depending on Birds have a four-chambered heart with
circulation designed for flight: Respiration is
 75

efficient, with air sacs and unidirectional flow to development of hair, and milk-producing
facilitate flight. Birds have a specialised excretory mammary glands. Initially young mammals are
system with no bladder; urine and faeces are completely dependent on milk from their mothers.
combined to produces “solid” or concentrated uric Milk is highly nutritious food which enables babies
acid. Marine birds have salt glands in their heads to to grow and develop rapidly. During this early phase
excrete excess salt. Paired ovaries and kidneys are of rapid growth, they have a set of “milk” teeth,
reduced to a single functional (left) ovary and which are later replaced by adult dentition, like back
kidney, which reduces the body weight of these teeth, or molars, which only erupt in this second set.
animals. Their nervous system is also specialised
Characteristics of mammals include hair or fur
nervous: the sensory system includes eyes with
for insulation, concealment (camouflage), signalling,
pectin (vascular organ attached to retina), and a
and waterproofing. Whiskers (or vibrissae) on
cochlea for hearing. Navigation takes place via
snouts, that functions as tactile organs or to signal
celestial bodies and magnetic fields.
mood in social mammals. Most mammalian orders
Class Mammalia (mammals) are placental mammals, with a long gestation and

Long before the evolution of dinosaurs and birds, well-developed babies when born, which then are

there existed a group of reptiles that had some nourished with highly nutritious and rich milk from

mammalian features. These mammal-like reptiles the female’s mammary glands. Mammals have large

occurred worldwide, but one of the richest sources brains with a neocortex (covering the cerebrum).

of their fossils is here in the Karoo, where fossils They also have a set of middle ear bones for

have been found showing the sequence of transmitting sound and four types of teeth (incisors,

anatomical changes that occurred during the canines, premolars, molars) that allows for a diverse

evolution of mammals from reptiles. diet.

The important changes that occurred during this The modern mammals can be divided into two

transition from reptiles to mammals include a loss of subclasses, the egg-laying (Prototheria) and live

the sprawling posture of reptiles; the loss of all but bearing (Theria).

one bone in the lower jaw; the development of The earliest mammals had young that hatched
teeth that can grind and slice food; a palate to from a leathery cleidoic egg. Indeed, the most
separate the mouth from the nasal passage (and thus primitive living mammals (Prototheria) still lay
enable chewing, suckling and breathing to occur eggs, but they are also endothermic, haired, and
simultaneously), and a larger brain. Many of these feed their young milk. These primitive mammals are
features are associated with the need to find and the monotremes (one order) with examples such as
rapidly process food to provide the energy to the duck-billed platypus and spiny anteaters from
regulate body temperature. Australia and New Guinea.

This is because mammals are “warm-blooded” More advanced living mammals are grouped
or endothermic. Other typically mammalian together as the Subclass Theria; all of them have a
features cannot be seen in fossils; these include the placenta that links the mother to the foetus
 76

developing within her uterus. Within the Theria two For perhaps a hundred million years, while the
groups can be distinguished: the Metatheria and dinosaurs ruled the earth, the mammals were
Eutheria. The Metatheria or marsupials include small, nocturnal, rather insignificant creatures. It
kangaroos, wallabies, possums, koalas. These was only after the extinction of the dinosaurs that
animals have poorly-developed placenta, a very they showed adaptive radiation into the niches left
short gestation length, and they give birth to very vacant by the dinosaurs. In time they became the
tiny, underdeveloped young that are (usually) dominant land vertebrates. Some species show
reared in a warm moist pouch in which the convergent evolution where members of different
mammary glands are situated. About 7 orders and animal lineages adapted to similar habitats on
250 species of marsupials occur today, also mostly in different continents by similar morphology. The
Australia and South America. The other group, the Eutheria have become the most abundant of all the
Eutheria, or placental mammals, have well mammals with about 3800 species living today; over
developed placentas, longer pregnancies and lack a a third of these are insectivores and bats.
pouch. The degree of development at birth varies a
There are 21 orders of living eutherian mammals,
lot and some new-born eutherian mammals are
classified largely by using features of the skull and
helpless (cats, dogs and humans are examples) while
teeth. Some of these orders have very few living
others are up and running within minutes of their
representatives while others are widely distributed
birth (all the antelope for example).
throughout the world. Two of the smaller orders that
These differences in the reproductive patterns of occur in southern Africa are of particular interest:
Metatheria and Eutheria reflect different the Hyracoidea (hyrax or dassie), and the
evolutionary responses to the stresses of the Tubulidentata (aardvark). Dassies display a curious
environment. The eutherians produce advanced mix of features and, surprisingly, seem to have their
young with superior chances of survival, but at a closest ancestral links with Perissodactyla (horses,
high and prolonged cost to the mother. The female rhinos), Probiscoidea (elephants), and the Sirenia
marsupial invests less energy before the birth of (manatees, sea cows, dugongs). The ancestry of
her young and can easily abandon her pouched aardvarks is even more uncertain; it has peg-like
babies, should environmental conditions become teeth, which have a structure quite different from
harsh; she can also rapidly produce more babies that found in other mammals. Aardvarks only occur
once conditions improve. In both groups the period in Africa south of the Sahara, and they live
of dependency on their mother provides an exclusively on termites that are eaten with a very
opportunity for learning, something particularly long tongue.
important in mammals with a long childhood (such
Most of the larger orders of mammals have
as many carnivores, elephants and primates) where
representatives in southern Africa. However, within
strong social bonding and exchange of information
these orders there are often interesting distribution
often occurs within the family.
patterns. For example, although the Artiodactyla
(even-toed ungulates) have a worldwide
 77

distribution, the sub-order that includes the deer torpid. At nightfall they warm up again, by
(they shed their horns each year) is totally absent metabolising fat, and then go and hunt for food. The
from Africa and is replaced by numerous species of camel in the desert needs to conserve both energy
antelope (buck). A similar situation is found in the and water. To do this the camel lets its temperature
Primata. The monkeys of Africa (Old World drop during the cold desert night until, eventually, a
monkeys) belong to a different group from those of body temperature of 34.5 °C is reached. The camel
the American continent (New World monkeys). then begins to shiver and uses energy to maintain its
temperature at that level. After daybreak, the camel
Several groups of mammals have returned to the
allows the sun to warm it up, this takes a long time
sea, from land-dwelling ancestors. Among these, the
because it is a large animal. If the day is very hot, an
Cetacea (whales and dolphins) are the only
upper body temperature of 40.5 °C is eventually
mammals, which never come onto land. The
reached and only then does the camel have to cool
prolonged and deep diving that the Cetacea
itself by evaporative water loss (sweating). By
undertake calls for extreme anatomical and
allowing these fluctuations in body temperature, a
physiological adaptations. The Cetacea have
camel can reduce its total daily water loss by half.
extremely large brains and are among the most
The gemsbok employs similar mechanisms to the
“intelligent” of all animals.
camel but, in addition, has evolved ways of keeping
Mammals, like the birds (and possibly some of the its brain cool and undamaged while its body
dinosaurs) show considerable control over their temperature rises to
body temperatures: They are endothermic (meaning 45 °C, a temperature that would otherwise be lethal.
that the body heat is produced from within the Many mammals living in very cold regions rely on
animal). Other vertebrates are ectothermic, and thick insulation (of dense fur or a thick layer of fat)
depend on environmental sources of heat to reach to reduce heat loss to the environment and thus
their preferred body temperature. Endothermy is a conserve energy.
superb way to become relatively independent of
Linked to their high-energy demands, mammals
many of the stresses of the physical environment,
(and birds) need a rapid delivery of oxygen to the
but it is only achieved through the expenditure of a
tissues. They have a complete, double blood
lot of energy. Endotherms need to eat a lot more
circulation in which the right side of the heart
food than do ectotherms.
receives deoxygenated blood from the body and
Many mammals (including humans) maintain an pumps it to the lungs to be oxygenated. The
almost constant body temperature. Some mammals, oxygenated blood is returned to the left side of the
however, allow their body temperature to rise and heart and a very muscular left ventricle then pumps
fall quite considerably. Thus, many small insect it, at high pressure, to the tissues and organs.
eating mammals (such as many bats) conserve
The chief nitrogenous excretory product in
energy by reducing their internal heat production
mammals is urea, a soluble end product that
during the daytime when they are not active; their
necessitates the loss of considerable amounts of
body temperature then drops, and they become
 78

water in the urine. It may at first seem odd that 5. Large brain, behaviour, and culture
mammals do not excrete dry uric acid as reptiles (transmission of learned behaviour between
and birds do. The answer lies in the fact that, while individuals and generations).
uric acid is the ideal excretory product for a
cleidoic egg, it is most unsuitable for an embryo
developing inside the mother and passing its waste
products into her blood stream via the placenta. In
this situation, the waste substances must be soluble
- hence the excretion of urea. Mammals do however
have some control of the amount of water they
excrete, and they have a special region in their
kidneys, the Loop of Henle, which enables them to
concentrate their urine. Indeed, desert mammals
(including the gemsbok) are so good at this that
they do not need to drink water but rely on
obtaining water from their food (often dry grasses
or seeds).

They have also modified their behaviour so that they

are active and feed at night when conditions are
most favourable to them.

Mammalian behaviour is fascinating and is often

complex. On the whole, probably because of their
relatively large brains, mammals place less reliance
on built-in instinctive behaviour patterns and have a
greater ability to learn from experience, being able
to adapt their behaviour according to circumstances
to an extent unrivalled by any other group of
animals. This reaches its peak in the primates.

Five trends led to uniquely human traits:

1. Enhanced daytime vision (binocular vision)

2. Upright (bipedal) walking: The location of
the foramen magnum is a key feature
separating four-legged and upright walkers
3. Better, more powerful grip, and precision
grip
4. Modified jaws and teeth (omnivorous diet)